Oviraptorosauria Barsbold, 1976
Definition- (Oviraptor philoceratops <- Therizinosaurus cheloniformis, Passer domesticus) (Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019; modified from Hu, Hou, Zhang and Xu, 2009)
Other definitions- (Oviraptor philoceratops <- Passer domesticus) (Maryanska, Osmólska and Wolsan, 2002; modified from Currie and Padian in Barsbold, 1997)
(Oviraptor philoceratops + Chirostenotes pergracilis) (modified from Padian, Hutchinson and Holtz, 1999)
(Oviraptor philoceratops <- Tyrannosaurus rex, Ornithomimus edmontonicus, Therizinosaurus cheloniformis, Troodon formosus, Passer domesticus) (Sereno, online 2005)
= Caenagnathiformes Sternberg, 1940
Definition- (Caenagnathus collinsi <- Passer domesticus) (Martyniuk, 2012)
= Avimimiformes Chatterjee, 1991
= Oviraptorosauria sensu Currie and Padian in Barsbold, 1997
Definition- (Oviraptor philoceratops <- Passer domesticus) (modified)
= "Oviraptoriformes" Sereno, online 2005
Definition- (Oviraptor philoceratops <- Passer domesticus) (Sereno, online 2005)
= Oviraptorosauria sensu Sereno, online 2005
Definition- (Oviraptor philoceratops <- Tyrannosaurus rex, Ornithomimus edmontonicus, Therizinosaurus cheloniformis, Troodon formosus, Passer domesticus)
= Oviraptoriformes Benson, 2008
Comments- Osborn (1924) described Oviraptor as an ornithomimid based on the toothless jaws, while Caenagnathus was originally (Sternberg, 1940) assigned to its own order Caenagnathiformes believed to be closer to crown Aves than hesperornithines or Ichthyornis. Cracraft (1971) further placed caenagnathiforms closest to galloanserines based on the sliding mandibular glenoid. Osmólska (1976) first noticed the similarity between her 'Oviraptor sp.' (actually Conchoraptor) and Caenagnathus, assigning both to Caenagnathidae within Theropoda. "The caenagnathids may represent a Cretaceous continuation of the same dinosaur lineage that gave rise to the birds", she said, and "a new suprafamilial taxon will have to be established as soon as the new material of Oviraptor has been described." This was done that same year when Barsbold (1976) erected Oviraptorosauria (incorrectly attributed in his paper to Barsbold, 1974) for oviraptorids. He wrote "the difference in the type of [edentulous] structure itself and undoubtedly the ecology of the oviraptorids and the ornithomimids are so great that their separation is presently fully valid." Barsbold viewed oviraptorosaurs as potentially diverging early from 'coelurosaurs' (coelophysoids and coelurid/compsognathid-grade taxa) due to supposedly primitive characters like the 'interclavicle' (actually a furcula unrecognized in other theropods at the time). By 1981 Barsbold had read Osmólska's paper and agreed with her the genera were closely related but wrote "considerable differences in the structure of the lower jaw of Caenagnathus and Oviraptor make it necessary to assign them to different families, which enter the infra-order Oviraptorosauria." Currie and Russell (1988) were the first authors to recognize Microvenator and the 'elmisaurids' (caenagnathid manus and pedes) belonged to Oviraptorosauria, but in 1994 Russell and Dong tried to expand the concept to include therizinosaurs, troodontids and ornithomimosaurs as well. Besides a few suggestions therizinosaurs might belong, this more inclusive Oviraptorosauria was not followed, and with the addition of basal taxa like Caudipteryx the group's content remains similar to this day. Note references to an "Oviraptorosauridae" as in Norell et al. (2000) are in error as there is no genus "Oviraptorosaurus" to base it on.
Avimimus was described in 1981 as closer to Aves than other theropods known at the time, then in the 1990s was associated with arctometatarsalian coelurosaurs. Currie (1989) was prescient in saying "similarities in the foot suggest that Avimimus may have been derived from elmisaurid dinosaurs" but Avimimus was first assigned to Oviraptorosauria in the first TWiG analysis (Norell et al., 2001). Chatterjee (1991) created a classification for Mesozoic birds where "each Mesozoic avian genus has been assigned to a corresponding higher level taxon or order" and so proposed an Avimimiformes for Avimimus, which has been ignored since then for being monotypic.
Sereno (online 2005) proposed the name Oviraptoriformes "to accommodate clades most closely related to Oviraptorosauria", with the phylogenetic definition "the most inclusive clade containing Oviraptor philoceratops Osborn 1924 but not Passer domesticus (Linnaeus 1758)." Yet online citations do not count for nomenclature. Benson (2008) is the first author to publish the name Oviraptoriformes, though he did such undefined in a phylogram without comment or definition. Holtz (2012) stated "Therizinosauria and Oviraptorosauria may together form a clade Oviraptoriformes" but again provided no definition. Martyniuk (2012) created a stem-based definition for Sternberg's (1940) Caenagnathiformes, generally unused since the 1970s, as all taxa closer to Caenagnathus collinsi than to Passer domesticus, which seemed more like an attempt to replace the newer Oviraptorosauria since he placed therizinosaurs further from birds. Yet it would work for a topology in which therizinosaurs were closer to oviraptorosaurs than to birds, is published with a proper definition and is an older name than Oviraptoriformes.
Paul (2016) used the informal term oviraptorosauriforms for "omnivoropterygids, oviraptorosaurs, and avians and their common ancestor, operative only if three groups form a clade that excludes all other dinosaurs." This would imply an 'Oviraptorosauriformes' with a partial definition (Omnivoropteryx sinousaorum + Oviraptor philoceratops + Passer domesticus, - XXX), but such a clade is poorly supported, taking 37 extra steps in Hartman et al.'s TWiG maniraptoromorph analysis for instance.
Oviraptorosauria defined- The first proposed phylogenetic definition of Oviraptorosauria was written in "A Note Added by the Editors" in Barsbold's (1997) encyclopedia entry, so is properly cited Currie and Padian in Barsbold, 1997. Their definition was "Oviraptoridae and all taxa closer to Oviraptor than to birds" which would have included therizinosaurs in most topologies over the following decade. Padian et al. (1999) noted this was inconsistant with common usage and so proposed a node-based definition instead- "Oviraptor and Chirostenotes (=Caenagnathus) and all descendants of their most recent common ancestor." However in the early 2000s taxa like Caudipteryx and Incisivosaurus were consistantly recognized as related to but outside this node and called oviraptorosaurs, so Hu et al. (2009) finally proposed a stem-based definition excluding therizinosaurs.
Ex-oviraptorosaurs- A partial sacrum and ilium (SMNS 58023) described by Frey and Martill (1995) as a possible oviraptorosaur, but was reidentified as a megaraptoran (Aranciaga Rolando et al., 2018).
A femur (ZIN PH 1/13) identified as oviraptorosaurian or ornithomimid by Nessov (1995) is ornithomimid (Averianov et al., 2003).
Currie et al. (1996) identified a surangular (NMV P186386) and dorsal (NMV P186302) from the Eumeralla Formation of Victoria as oviraptorosaurian, but these have been placed more ambiguously as theropod (Agnolin et al., 2010) and maniraptoran (Benson et al., 2012) lately.
A cervical vertebra (MACN 622) discovered with the Noasaurus holotype was originally identified as an oviraptorosaur (Frankfurt and Chiappe, 1999), but reidentified by Agnolin and Martinelli (2007) as a noasaurid, and probably part of the holotype individual.
Paul (2010) placed Sapeornis in Oviraptorosauria, but that takes 37 more steps in Hartman et al.'s (2019) matrix so is highly unlikely. Paul also placed Epidexipteryx there (but not Scansoriopteryx), and Brusatte et al. (2014) recovered an Epidexipteryx plus Pedopenna pair as the basalmost oviraptorosaurs. Cau (2018) recovered all scansoriopterygids as oviraptorosaurs in his Bayesian analysis. In a modified version of the Hartman et al. matrix, scansoriopterygids move to Oviraptorosauria in 9 steps, but getting Epidexipteryx or Pedopenna there without Scansoriopteryx takes 13 steps and 4 steps respectively.
References- Osborn, 1924. Three new Theropoda, Protoceratops zone, central Mongolia. American Museum Novitates. 144, 1-12.
Sternberg, 1940. A toothless bird from the Cretaceous of Alberta. Journal of Paleontology. 14(1), 81-85.
Cracraft, 1971. Caenagnathiformes: Cretaceous birds convergent in jaw mechanism to dicynodont reptiles. Journal of Paleontology. 45(5), 805-809.
Barsbold, 1976. The evolution and systematics of late Mesozoic carnivorous dinosaurs. In Kramarenko, Luvsandansan, Voronin, Barsbold, Rozhdestvensky, Trofimov and Reshetov (Eds.). Paleontology and Biostratigraphy of Mongolia. The Joint Soviet-Mongolian Paleontological Expedition, Transactions. 3, 68-75.
Osmólska, 1976. New light on skull anatomy and systematic position of Oviraptor. Nature. 262, 683-684.
Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological Expedition Transactions. 15, 28-39.
Currie and Russell, 1988. Osteology and relationships of Chirostenotes pergracilis (Saurischia, Theropoda) from the Judith River (Oldman) Formation of Alberta, Canada. Canadian Journal of Earth Sciences. 25(7), 972-986.
Currie, 1989. Theropod dinosaurs of the Cretaceous. In Padian and Chure (Eds.). The Age of Dinosaurs. Short Courses in Paleontology. 2, 113-120.
Chatterjee, 1991. Cranial anatomy and relationships of a new Triassic bird from Texas. Philosophical Transactions of the Royal Society of London Series B. 332(1265), 277-342.
Russell and Dong, 1994. The affinities of a new theropod from the Alxa Desert, Inner Mongolia, People’s Republic of China. Canadian Journal of Earth Sciences. 30(10), 2107-2127.
Frey and Martill, 1995. A possible oviraptorosaurid theropod from the Santana Formation (Lower Cretaceous, Albian?) of Brazil. Neues Jahrbuch Fur Geologie und Palaeontologie. 7, 397-412.
Nessov, 1995. Dinosaurs of nothern Eurasia: new data about assemblages, ecology, and paleobiogeography. Institute for Scientific Research on the Earth's Crust, St. Petersburg State University, St. Petersburg. 1-156.
Currie, Vickers-Rich and Rich, 1996. Possible oviraptorosaur (Theropoda, Dinosauria) specimens from the Early Cretaceous Otway Group of Dinosaur Cove, Australia. Alcheringa. 20(1-2), 73-79.
Barsbold, 1997. Oviraptorosauria. In Currie and Padian (eds.). Encyclopedia of Dinosaurs. 505-509.
Frankfurt and Chiappe, 1999. A possible oviraptorosaur from the Late Cretaceous of northwestern Argentina. Journal of Vertebrate Paleontology. 19(1), 101-105.
Padian, Hutchinson and Holtz, 1999. Phylogenetic definitions and nomenclature of the major taxonomic categories of the carnivorous Dinosauria (Theropoda). Journal of Vertebrate Paleontology. 19(1), 69-80.
Norell, Makovicky and Clark, 2000. A new troodontid theropod from Ukhaa Tolgod, Mongolia. Journal of Vertebrate Paleontology. 20(1), 7-11.
Norell, Clark and Makovicky, 2001. Phylogenetic relationships among coelurosaurian theropods. In Gauthier and Gall (eds.). New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom. 49-67.
Maryanska, Osmólska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47(1), 97-116.
Averianov, Starkov and Skutschas, 2003. Dinosaurs from the Early Cretaceous Murtoi Formation in Buryatia, eastern Russia. Journal of Vertebrate Paleontology. 23(3), 586-594.
Sereno, online 2005. Stem Archosauria - TaxonSearch. http://www.taxonsearch.org/dev/file_home.php [version 1.0, 2005 November 7]
Agnolin and Martinelli, 2007. Did oviraptorosaurs (Dinosauria; Theropoda) inhabit Argentina? Cretaceous Research. 28(5), 785-790.
Balanoff, Bever and Rowe, 2008. The endocranial morphology of oviraptorosaurs and a reinterpretation of their encephalization quotients. Journal of Vertebrate Paleontology. 28(3), 47A.
Benson, 2008. New information on Stokesosaurus, a tyrannosauroid (Dinosauria: Theropoda) from North America and the United Kingdom. Journal of Vertebrate Paleontology. 28(3), 732-750.
Hu, Hou, Zhang and Xu, 2009. A pre-Archaeopteryx troodontid theropod from China with long feathers on the metatarsus. Nature. 461, 640-643.
Agnolin, Ezcurra, Pais and Salisbury, 2010. A reappraisal of the Cretaceous non-avian dinosaur faunas from Australia and New Zealand: Evidence for their Gondwanan affinities. Journal of Systematic Palaeontology. 8(2), 257-300.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press. 320 pp.
Balanoff, 2011. Oviraptorosauria: Morphology, phylogeny, and endocranial evolution. PhD thesis. Columbia University. 522 pp.
Persons, Currie and Norell, 2011. Shake your feathers: The flamboyant, athletic, and possibly flirtatious caudal morphology of oviraptorosaurs. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 174.
Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna from southern Australia indicates high polor diversity and climate-driven dinosaur provinciality. PLOS One. 7(5), e37122.
Holtz, 2012. Theropods. In Brett-Surman, Holtz and Farlow (Eds.). The Complete Dinosaur. Second edition. Indiana University Press. 347-378.
Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs. Vernon, New Jersey. Pan Aves. 189 pp.
Balanoff, Bever and Norell, 2013. The relationships of oviraptorosaurian dinosaurs and endocranial evolution along a morphologically bizarre lineage. Journal of Vertebrate Paleontology. Program and Abstracts 2013, 81.
Pittman and Hutchinson, 2013. The evolution of tail joint stiffness in oviraptorosaur dinosaurs and its consequences for tail function. Journal of Vertebrate Paleontology. Program and Abstracts 2013, 191.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan culminated in rapid rates of evolution across the dinosaur-bird transition. Current Biology. 24(20), 2386-2392.
Tanaka, Zelenitsky, Lu, Yi, Pu, Chang, Xu and Li, 2014. Nest type and incubation behavior in oviraptorosaurs in relation to body size. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 238-239.
Paul, 2016. The Princeton Field Guide to Dinosaurs 2nd edition. Princeton University Press. 360 pp.
Cau, 2018. The assembly of the avian body plan: A 160-million-year long process. Bollettino della Società Paleontologica Italiana. 57(1), 1-25.
Aranciaga Rolando, Brisson Egli, Sales, Martinelli, Canale and Ezcurra, 2018 (online 2017). A supposed Gondwanan oviraptorosaur from the Albian of Brazil represents the oldest South American megaraptoran. Cretaceous Research. 84, 107-119.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

undescribed oviraptorosaur (Currie, 2002)
Aptian-Albian, Early Cretaceous
Ohshih Formation, Mongolia
Material- (PJC.2001.10) proximal femur
Reference- Currie, 2002. Report on fieldwork in Mongolia, September 2001. Alberta Palaeontological Society, sixth annual symposium. 8-12.

undescribed Oviraptorosauria (Watabe and Suzuki, 2000)
Late Campanian, Late Cretaceous
Dzamin Khond, Djadochta Formation, Mongolia
Material- elements
Comments- Watabe and Suzuki (2000) state "isolated bones of oviraptorosauria" were found in 1993, 1994 and/or 1998.
Reference- Watabe and Suzuki, 2000. Cretaceous fossil localities and a list of fossils collected by the Hayashibara Museum of Natural Sciences and Mongolian Paleontological Center Joint Paleontological Expedition (JMJPE) from 1993 through 1998. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 99-108.

undescribed Oviraptorosauria (Suzuki and Watabe, 2000)
Early Maastrichtian, Late Cretaceous
Altan Uul, Nemegt Formation, Mongolia
Material- ?(IGM coll.; 060908 AU-IV BON) partial skeleton including pelvic elements (Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010)
Early Maastrichtian, Late Cretaceous
Bugin Tsav, Nemegt Formation, Mongolia
(IGM coll.; 980803 BgT NAR) dorsal vertebrae, caudal vertebrae, hindlimb elements (Suzuki and Watabe, 2000)
(IGM coll.; 060811 BgT-II KHTB) partial skeleton (Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010)
Early Maastrichtian, Late Cretaceous
Gurilin Tsav, Nemegt Formation, Mongolia
(uncollected?) elements
Early Maastrichtian?, Late Cretaceous
Khermeen Tsav, Nemegt Formation?, Mongolia
(IGM coll.; 060904 KmT KHTB Ovi) partial skull, dorsal vertebrae, sternal, forelimb, pelvis, hindlimb (Matsumoto, Hashimoto, Sonoda, Fujiyama, Mifune, Kawahara and Saneyoshi, 2010)
Early Maastrichtian, Late Cretaceous
Upper White Beds of Khermeen Tsav, Nemegt Formation, Mongolia
(IGM coll.; 060906 KmT KHTB) (four individuals)
Early Maastrichtian, Late Cretaceous
Tsagaan Khushuu, Nemegt Formation, Mongolia
(uncollected?) elements (Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010)
Comments- These are field numbers for a joint HMNS-IGM collection.
980803 BgT NAR was found in 1994 at Bugin Tsav with a small Tarbosaurus specimen, preserves cervicodorsal hypapophyses and was at the IGM as of 2000 (Suzuki and Watabe, 2000).
Matsumoto et al. (2010) list 060904 KmT KHTB Ovi as Oviraptorosauria found on September 4 2004, but this would indicate the field number would be 040904 and be from Cenozoic locality Baga Dzan. It's more likely this was found on September 4 2006. Prepared contents are listed as "dorsal vertebrae, pelvic part and hindlimb" and "part of skull, forelimb and sternal."
Watabe et al. (2010) wrote "isolated bones and partial skeletons of Oviraptorosauria (Theropoda)" were discovered in 2006 at Bugin Tsav, of which 060811 BgT-II KHTB was labeled Oviraptorosauria and found on June 8. They also stated "isolated bones of Oviraptorosauria" were found in 2006 at Gurilin Tsav but did not list collected specimens. "A partial skeleton
of small theropod" was said to be found that year at Altan Uul, which may correspond to 060908 AU-IV BON discovered on September 8 at Altan Uul IV, labeled as "?Oviraptorosauria" with material listed as "pelvic." Watabe et al. say "isolated and articulated bones of ... Oviraptorosauria" were found at Tsaagan Khushuu in 2006, but did not list collected material. They stated "the associated bones of Oviraptorosauria including cranial elements were found" in the Upper White Beds of Khermeen Tsav in 2006, and that "those bones are from more than four individuals of mass-burial in a single layer." Watabe et al.'s list indicates the specimen number 060906 KmT KHTB, found on September 6.
References- Suzuki and Watabe, 2000. Report on the Japan - Mongolia Joint Paleontological Expedition to the Gobi desert, 1998. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 83-98.
Matsumoto, Hashimoto, Sonoda, Fujiyama, Mifune, Kawahara and Saneyoshi, 2010. Report of the preparation works for Mongolian specimens in Hayashibara Museum of Natural Sciences: 1999-2008. Hayashibara Museum of Natural Sciences Research Bulletin. 3, 167-185.
Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010. Report of the HMNS-MPC Joint Paleontological Expedition in 2006. Hayashibara Museum of Natural Sciences Research Bulletin. 3, 11-18.

Chuniaoia Ji and Ji, 2001
Comments- Ji and Ji (2001) used the name Chuniaoia in a cladogram for a branch leading to Protarchaeopteryx, but it has not been defined nor has its intended purpose been published. It is similar to the name Chuniaoae used in the online supplementary information of Ji et al. (1998), which was seemingly for a Caudipteryx+Aves clade.
Ji et al. (1998) gave the title of Protarchaeopteryx's description as "Protarchaeopterygid bird (Protarchaeopteryx gen. nov.) - fossil remains of archaeopterygids from China", while Downs' 2001 translation is titled "A Chinese archaeopterygian, Protarchaeopteryx gen. nov.". The first four characters (恩错祖鸟) translate to Protarchaeopteryx, so "protarchaeopterygid" is an invention of Ji et al., made more obvious by the fact it was classified as an archaeopterygid in the 1997 paper. The implied family 'Protarchaeopterygidae' has never been published so remains unofficial. Google Translate gives the translation "Protarchaeopteryx gen. nov. - Archaeopteryx of China."
References- Ji and Ji, 1997. 恩错祖鸟(Protarchaeopteryx gen. nov.) - 中国的始祖鸟类亿右. Chinese Geology. 24(3) (total issue 238), 38-41, 49.
Ji, Currie, Norell and Ji, 1998. Two feathered dinosaurs from northeastern China. Nature. 393, 753-761.
Ji and Ji, 2001. How can we define a feathered dinosaur as a bird? In Gauthier and Gall (eds.). New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom. 43-46.
Protarchaeopteryx Ji and Ji, 1997
P. robusta Ji and Ji, 1997
Late Barremian-Early Aptian, Early Cretaceous
Jianshangou member of Yixian Formation, Liaoning, China
Holotype- (NGMC 2125) (690 mm) incomplete skull (70 mm), mandibles, hyoid, six cervical vertebrae (16 mm), five dorsal vertebrae (13 mm), sacrum, fourteen caudal vertebrae, chevron, proximal scapula, partial coracoids, sternal plates (25x15 mm), partial furcula, partial humeri (87 mm), incomplete radii (73 mm), incomplete ulnae (74 mm), scapholunare, semilunate carpal, distal carpal III, metacarpal I (17 mm), phalanx I-1, manual ungual I, metacarpal II (44 mm), phalanx II-1, phalanx II-2, manual ungual II, metacarpal III (45 mm), phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, two keratinous manual ungual sheaths, partial ilium (95 mm), incomplete pubes (80 mm), ischial fragment(?), incomplete femora (125 mm), tibiae (160 mm), partial fibulae, astragali, calcaneum, distal tarsal III, distal tarsal IV, metatarsal I (13 mm), phalanx I-1, pedal ungual I, metatarsal II (77 mm), phalanx II-1, phalanx II-2, pedal ungual III, metatarsal III (86 mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV (81 mm), phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal V, contour feathers, retrices
Diagnosis- (after Senter et al., 2004) six maxillary teeth; seven dentary teeth.
Other diagnoses- Ji and Ji's (1997) initial diagnosis is formed entirely of plesiomorphies (claviform and unserrated dentition; sternum is thin and flat; tail is long; forelimb/hindlimb index is 0.7; forelimb resembles Archaeopteryx in morphology with three unguals, the second of which is enlarged; ilium is large and elongated; pubes are robust and distally fused; hind limb is long and robust with digit I reduced and dorsally migrated; torso feathers are 50 mm in length with short and robust shafts; tail fan is extremely well developed; retrices are as long as 150 mm with a slender and elongated shaft and slender and gracile barbs) and untrue statements (proximal metatarsals are fused; pedal digit I lies in opposition to digit III and forms a grasping apparatus).
Ji et al. (1998) also included the following characters in their diagnosis- large straight premaxillary teeth (also in Incisivosaurus); short, bulbous maxillary and dentary teeth (plesiomorphic for Maniraptora); all of which are primitively serrated (incorrect- Senter et al., 2004); rectrices form a fan at the end of the tail (plesiomorphic for Pennaraptora).
Comments- The holotype was discovered in 1996.
Phylogenetic relationships- Ji and Ji (1997) initially referred Protarchaeopteryx to Archaeopterygidae based on constricted tooth roots ("claviform" teeth; primitive for maniraptorans) and "elongated forelimb or primitive wing with three talons" (primitive for maniraptoriforms) and "extremely well developed tail fan, and feather morphology" (primitive for pennaraptorans).
Zhou (1997) classified Protarchaeopteryx as a sauriurine bird more closely related to enantiornithines than Confuciusornis due to the smaller manual ungual III, but Sauriurae is near universally rejected and subsequent analyses have shown Confuciusornis is far more similar to enantiornithines than Protarchaeopteryx.
Xu et al. (1999) resolved Protarchaeopteryx as a paravian in a trichotomy with Caudipteryx and Troodontidae+Eumaniraptora. Xu et al. (2000) used 86 of the same characters plus three new ones, added Microraptor and removed Tyrannosauridae and Unenlagia. Their tree was slightly more resolved, as Protarchaeopteryx was found to be the sister taxon of Troodontidae+Eumaniraptora, and thus closer to birds than Caudipteryx. However, both studies supported the paravian status of Protarchaeopteryx using the same seven characters. Of these, three (deep suborbital bar; unfused interdental plates; pubic foot projects posteriorly only) are unknown in Protarchaeopteryx, and two others (less than 11 caudal vertebrae with transverse processes; metacarpal I less than 33% of metacarpal II in length) are not present in the taxon. Oviraptorids and troodontids are miscoded as lacking a proximodistally elongate coracoid, which is actually a pennaraptoran synapomorphy. A radius less than 70% of ulnar width is shared with e.g. Caudipteryx, Similicaudipteryx and Anzu and is highly homoplasious in basal Paraves.
Xu et al. (2000) resolved Protarchaeopteryx as a paravian more derived than Caudipteryx based on four additional characters. Two (teeth with serrated mesial and distal carinae; mid and distal caudal vertebrae at least 130% longer than proximal caudals) arn't present in Protarchaeopteryx , while a short dorsal process of the premaxilla cannot be determined. The remaining character (manus over 120% of ulnar length) was misscored as lacking in Compsognathus, ornithomimids, Caudipteryx and oviraptorids (ornithomimids and oviraptorids are polymorphic), but is also a composite character involving lengths of different manual elements.
Holtz (2001) found Protarchaeopteryx to be either a basal member of the therizinosaur-oviraptorosaur clade, sister taxon to Pennaraptora, or a basal paravian.
Ji and Ji (2001) placed Protarchaeopteryx as a basal avialan in their cladogram, using the name Dromavialae for the node of Protarchaeopteryx+Aves. This was based on the character "real wings with symmetrical feathers of modern concept", which is vague and also occurs in basal pennaraptorans.
Gishlick (2002) found Protarchaeopteryx to be in a eumaniraptoran polytomy with Deinonychus, Sinornithosaurus, Archaeopteryx and Pygostylia, though the matrix only included forelimb characters.
Paul (2002) assigned Protarchaeopteryx to the Archaeopterygidae based on several characters, none of which are convincing. For instance, the teeth are virtually identical to those of Incisivosaurus, so Protarchaeopteryx's "small, conical teeth" are not distinctively archaeopterygid. The sternal morphology is invalid because Archaeopteryx's supposed sternum is actually a coracoid. Other characters such as the "slender fingers", and "non-fused but tighly articulated metacarpals, tarsals and metatarsals" are symplesiomorphic within maniraptorans and vague. Only one third manual digit of Protarchaeopteryx is crossed under digit II, which is taphonomic instead of structural in any case. The ilium is too incomplete to know if the preacetabular process would make it parallelogram-shaped, while the pubis is too incomplete to measure pelvic canal depth. I cannot confirm the coracoid is strongly bent as in avepectorans. The robust second manual digit is shared with caenagnathids, while the low ilium and slender pointed postacetabular process are unquantified but also present in e.g. Similicaudipteryx.
Senter (2003) combined Protarchaeopteryx and Incisivosaurus into one OTU, which he found to be the basalmost oviraptorosaur. Senter et al. (2004) came to the same conclusion.
Holtz et al. (2004) recovered Protarchaeopteryx as a paravian more derived than alvarezsaurids, excluded from Deinonychosauria and Archaeopteryx+Ornithurae.
Senter (2007) and Cau (2018) found Protarchaeopteryx to be the sister group of Incisivosaurus, with both of these taxa as the most basal oviraptorosaurs.
Funston and Currie (2016) added the taxon to a version of Maryanska's oviraptorosaur matrix and found it emerged as the sistergroup of Archaeopteryx, albeit with Velociraptor and Herrerasaurus as the only other included non-oviraptorosaurs.
Hartman et al. (2019) recovered Protarchaeopteryx as the sister taxon to Pennaraptora, but after adding more taxa it is the most basal oviraptorosaur. In that updated matrix it takes 13 steps to be an archaeopterygid, 34 steps to be between Confuciusornis and enantiornithines, 5 steps to be a paravian,
References- Ji and Ji, 1997. 恩错祖鸟(Protarchaeopteryx gen. nov.) - 中国的始祖鸟类亿右. Chinese Geology. 24(3) (total issue 238), 38-41, 49.
Zhou, 1997. Diversification of birds from the "Late Jurassic" of China. Journal of Vertebrate Paleontology. 17(3), 86A.
Ji, Currie, Norell and Ji, 1998. Two feathered dinosaurs from northeastern China. Nature. 393, 753-761.
Xu, Wang and Wu, 1999. A dromaeosaurid dinosaur with filamentous integument from the Yixian Formation of China. Nature. 401, 262-266.
Xu, Zhou and Wang, 2000. The smallest known non-avian theropod dinosaur. Nature. 408, 705-708.
Holtz, 2001. Arctometatarsalia revisited: the problem of homplasy in reconstructing theropod phylogeny. pp. 99-122. in Gauthier and Gall (eds.). New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom. Yale Univ. Press.
Ji and Ji, 2001. How can we define a feathered dinosaur as a bird? In Gauthier and Gall (eds.). New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom. 43-46.
Padian, Ji and Ji, 2001. Feathered dinosaurs and the origin of flight. in Tanke and Carpenter (eds.). Mesozoic Vertebrate Life. 117-138.
Gishlick, 2002. The functional morphology of the forelimb of Deinonychus antirrhopus and its importance for the origin of avian flight. Unpublished PhD thesis. Yale University, 142 pp.
Paul, 2002. Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds. Baltimore: Johns Hopkins University Press.
Senter, 2003. Taxonomic sampling artifacts and the phylogenetic position of Aves. Unpublished PhD thesis. Northern Illonois University.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria. Second Edition. University of California Press. 71-110.
Senter, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae. Bulletin of Gunma Museum of Natural History 8: 1-20.
Senter, 2007. A new look at the phylogeny of Coelurosauria. Journal of Systematic Palaeontology. 5(4), 429-463.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.
Cau, 2018. The assembly of the avian body plan: A 160-million-year long process. Bollettino della Società Paleontologica Italiana. 57(1), 1-25.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247.

Similicaudipteryx He, Wang and Zhou, 2008
S. yixianensis He, Wang and Zhou, 2008
= Caudipteryx yixianensis (He, Wang and Zhou, 2008) Paul, 2010
Early Albian, Early Cretaceous
Jiufotang Formation, Liaoning, China
Holotype- (IVPP V12556) (adult) six cervical vertebrae, cervical ribs, fourteen dorsal vertebrae, dorsal rib fragments, sacrum (85 mm), twenty-one partial caudal vertebrae, pygostyle, six chevrons, incomplete scapula, partial coracoid, incomplete sternal plates, sternal rib fragments, humerus (~130 mm), partial ulna, partial radius, ilia (one partial; 153 mm), pubes (~223 mm), ischial fragment, femora (one distal; ~220 mm), tibiae (one incomplete; ~240 mm), partial fibula (~223 mm), metatarsal I (25 mm), phalanx I-1 (28 mm), pedal ungual I (23 mm), metatarsals II (144 mm), phalanges II-1 (~43 mm), phalanx II-2 (39 mm), pedal ungual II (33 mm), metatarsals III (~183 mm), phalanges III-1 (46 mm), phalanx III-2 (34 mm), phalanx III-3 (32 mm), pedal ungual III (34 mm), metatarsals IV (153 mm), phalanges IV-1 (30 mm), phalanx IV-2 (23 mm), phalanx IV-3 (18 mm), phalanx IV-4 (21 mm), pedal ungual IV (20 mm)
Late Valanginian-Middle Aptian, Early Cretaceous
Yixian Formation, Liaoning, China
Referred- (STM4-1) (juvenile) partial skull, cervical vertebrae, dorsal vertebrae, dorsal ribs, gastralia, caudal vertebrae, chevrons, scapulae, humeri (24 mm), radii (21 mm), ulnae, semilunate carpals, metacarpals I, phalanx I-1, manual ungual I, metacarpals II, phalanges II-1, phalanges II-2, manual unguals II, metacarpals III, phalanges III-1, phalanges III-2, phalanx III-3, manual ungual III, ilium, pubis, ischium, femora (38 mm), tibiae (49 mm), fibula, astragalus, metatarsals I, phalanges I-1, pedal unguals I, metatarsals II (one incomplete), phalanges II-1 (one incomplete), partial phalanx II-2, metatarsals III (one incomplete), phalanges III-1 (one incomplete), phalanx III-2, phalanx III-3 fragment, metatarsals IV (one partial), phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, incomplete pedal ungual IV, metatarsal V, body feathers, remiges, retrices (Xu et al., 2010a)
(STM22-6) (subadult) skull, mandible, ten cervical vertebrae, cervical ribs, dorsal vertebrae, dorsal ribs, gastralia, caudal vertebrae, pygostyle, chevrons, scapulae, coracoid, furcula, humeri (81 mm), radii (69 mm), ulnae, metacarpals I, phalanges I-1, manual unguals I, metacarpals II, phalanges II-1, phalanx II-2, manual ungual II, metacarpals III, phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, incomplete ilium, pubes, ischium, femur (140 mm), tibiae (182 mm), astragali, phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, body feathers, remiges, retrices (Xu et al., 2010a)
Diagnosis- (modified from He et al., 2008) puboilial ratio of 1.46.
Other diagnoses- In its diagnosis, He et al. list a dagger-like pygostyle (also present in Nomingia and Citipati), ilium shaped like Caudipteryx (ambiguous), two large anterior dorsal hypapophyses (present in caenagnathoids) and puboilial ratio of 1.46. The latter compares with 1.04-1.12 in Caudipteryx, .96 in Nomingia and .99 in Microvenator.
Comments- The holotype was first mentioned by Wang et al. (2007) as an unnamed caudipterid and later described and named by He et al. (2008). Xu et al. (2010a) briefly describe two new, younger specimens which are almost complete.
He et al. referred it to Caudipteridae based on several characters. Most are symplesiomorphic for maniraptorans (low number of caudal vertebrae; deep pubic peduncle; unfused metatarsus; metatarsal III longest; metatarsals II and IV subequal in width; metatarsal II slightly shorter than IV; subarctometatarsal metatarsus), the proximally placed metatarsal I is also present in Protarchaeopteryx, and the preacetabular depth is intermediate between other oviraptorosaurs.
Xu et al. (2011) added Similicaudipteryx to Senter's TWiG matrix and recovered it as an oviraptorosaur more derived than Protarchaeopteryx.
Lamanna et al. (2014) added this to Maryanska's oviraptorosaur matrix and recovered it as a caudipterid.
Cau (2018) recovered it as more derived than Protarchaeopteryx and Incisivosaurus, but less than Caudipteryx.
Most recently, Hartman et al. (2019) recovered it as the most basal oviraptorosaur, but after adding several taxa Protarchaeopteryx moves to be the basalmost member instead.
References- Wang, Jones and Evans, 2007. A juvenile anuran from the Lower Cretaceous Jiufotang Formation, Liaoning, China. Cretaceous Research. 28, 235-244.
He, Wang and Zhou, 2008. A new genus and species of caudipterid dinosaur from the Lower Cretaceous Jiufotang Formation of Western Liaoning, China. Vertebrata PalAsiatica. 46(3), 178-189.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press. 320 pp.
Ptum, 2010. Moulting tail feathers in a juvenile oviraptorisaur. Nature. 468, E1.
Xu, Zheng and Yu, 2010a. Exceptional dinosaur fossils show ontogenetic development of early feathers. Nature. 464, 1338-1341.
Xu, Zheng and Yu, 2010b. Xu et al. reply. Nature. 464, 468, E2.
Xu, You, Du and Han, 2011. An Archaeopteryx-like theropod from China and the origin of Avialae. Nature. 475, 465-470.
Lamanna, Sues, Schachner and Lyson, 2014. A new large-bodied oviraptorosaurian theropod dinosaur from the Latest Cretaceous of Western North America. PLoS ONE. 9(3), e92022.
Cau, 2018. The assembly of the avian body plan: A 160-million-year long process. Bollettino della Società Paleontologica Italiana. 57(1), 1-25. DOI: 10.4435/BSPI.2018.01
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

Ganzhousaurus Wang, Sun, Sullivan and Xu, 2013
G. nankangensis Wang, Sun, Sullivan and Xu, 2013
Late Cretaceous
Nanxiong Group, near Ganzhou, Jiangxi, China
Holotype- (SDM 20090302) incomplete mandible, distal caudal vertebra, incomplete distal caudal vertebra, partial distal caudal vertebra, ilial fragment, partial tibia, distal tarsal III, metatarsal I (36 mm), phalanx I-1 (28 mm), pedal ungual I, metatarsal II (125 mm), phalanx II-1 (41 mm), phalanx II-2, partial pedal ungual II, metatarsal III (147 mm), phalanx III-1 (45 mm), partial phalanx III-2, phalanx IV-1, partial phalanx IV-2
Diagnosis- (after Wang et al., 2013) shallow dentary (ratio of maximum anteroposterior length to maximum dorsoventral depth 1.90); absence of fossa or foramen on lateral surface of dentary (also in oviraptorids; unlike Gigantoraptor, Chirostenotes and Caenagnathasia); weakly downturned anterior mandibular end (also in Khaan; unlike Nemegtomaia and Heyuannia); shallow depression immediately surrounding anterior margin of external mandibular fenestra (also in Gigantoraptor, Nemegtomaia and Heyuannia; unlike Citipati and Khaan); external mandibular fenestra subdivided by anterior process of surangular (also in oviraptorids); dentary posteroventral process slightly twisted and positioned on mandibular ventrolateral surface (also in Nemegtomaia; unlike Citipati and Heyuannia); shallow longitudinal groove along medial surface of dentary posteroventral process (also in Microvenator; unknown in other oviraptorosaurs); angular anterior process wider transversely than deep dorsoventrally (unknown in other oviraptorosaurs); sharp groove along ventrolateral surface of angular anterior process (also in Chirostenotes, Gigantoraptor and Khaan; unlike most oviraptorosaurs); ventral border of external mandibular fenestra formed mainly by angular (also in Chirostenotes and Gigantoraptor; unlike oviraptorids); ventral flange along distal half of metatarsal II (also in Avimimus; unlike Citipati, Wulatelong and Heyuannia); arctometatarsal condition absent (also in oviraptorids; unlike Avimimus, Chirostenotes and Elmisaurus).
Comments- The holotype was purchased from a fossil dealer and initially mentioned in an abstract by Wang and Xu (2012) before being officially described by Wang et al. (2013). Initially the partial tibia was thought to be a femur by Wang and Xu. Note the diagnosis given by Wang et al. includes numerous symplesiomorphies and will need to be revised.
Wang and Xu found it to be a heyuannine in their unpublished analysis, while Wang et al. found it to be most closely related to Citipati and Rinchenia in its published form, while Cau (online, 2013) found it to be in a trichotomy with caenagnathids and oviraptorids. Hartman et al. (2019) found it to be between Similicaudipteryx and caudipterids.
References- Wang and Xu, 2012. A new oviraptorid specimen from the Upper Cretaceous of southern China. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 190.
Cau, online 2013. http://theropoda.blogspot.com/2013/04/ganzhousaurus-nankangensis-wang-et-al.html
Wang, Sun, Sullivan and Xu, 2013. A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of southern China. Zootaxa. 3640(2), 242-257.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

undescribed oviraptorosaur (Zhou, O'Connor and Wang, 2014)
Early Albian, Early Cretaceous
Sihedang, Jiufotang Formation, Liaoning, China
Comments- Zhou et al. (2014) reported "one caudipteryid (Maniraptora: Oviraptorosauria)" from Sihedang. This may be "Lingyuanosaurus", which was described in 2019 as a therizinosaur, but has oviraptorosaurian characters as well.
Reference- Zhou, O'Connor and Wang, 2014. A new species from an ornithuromorph (Aves: Ornithothoraces) dominated locality of the Jehol Biota. Chinese Science Bulletin. 59(36), 5366-5378.

Caudipteridae Zhou and Wang, 2000
Definition- (Caudipteryx zoui <- Caenagnathus collinsi, Oviraptor philoceratops) (Hendrickx, Hartman and Mateus, 2015)
= Caudipterygidae Osmólska, Currie and Barsbold, 2004
Comments- In 2000, Zhou and Wang proposed the family Caudipteridae for Caudipteryx. Osmólska et al. (2004) emmended this to Caudipterygidae, since Caudipteridae is formed incorrectly (ICZN Article 29.3). It has been suggested this is unecessary, since according to Article 29.4, "if after 1999 a new family-group name is based on a generic name which is or ends in a Greek or Latin word or ends in a Greek or Latin suffix, but its derivation does not follow the grammatical procedures of Articles 29.3.1 or 29.3.2, its original spelling must be maintained as the correct original spelling." However, Article 29.4.2 states this is only true provided the genus was treated as an arbitrary combination of letters (e.g. "Caudipteryxidae"), which is not the case. To complicate matters, Article 29.5 states "If a spelling of a family-group name was not formed in accordance with Article 29.3 but is in prevailing usage, that spelling is to be maintained, whether or not it is the original spelling and whether or not its derivation from the name of the type genus is in accordance with the grammatical procedures in Articles 29.3.1 and 29.3.2." Caudipteridae has 6420 search results compared to Caudipterygidae's 402 on Google, and 15 vs. 14 on Google Scholar (as of 8-26-19). Thus Caudipteridae should be maintained.
References- Zhou and Wang, 2000. A new species of Caudipteryx from the Yixian Formation of Liaoning, northeast China. Vertebrata PalAsiatica. 38(2), 113-130.
Osmólska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel, Dodson and Osmólska, (eds.). The Dinosauria, Second Edition. University of California Press. 165-183.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.

Caudipteryx Ji, Currie, Norell and Ji, 1998
Diagnosis- premaxillary teeth limited to rostral half of element; first premaxillary tooth much larger than others (also in Incisivosaurus and Protarchaeopteryx); single maxillary fenestra present; twenty-two caudal vertebrae; sternal plates oval; only two phalanges present on manual digit III.
Other diagnoses- (after Ji et al., 1998) elongate, hooked premaxillary teeth with broad roots; maxilla and dentary edentulous; tail short (one-quarter of the length of the body); arm long for a non-avian theropod; short manual unguals; Leg-to-arm ratio, 2.5.
Reference- Ji, Currie, Norell and Ji, 1998. Two feathered dinosaurs from northeastern China. Nature. 393, 753-761.
C. zoui Ji, Currie, Norell and Ji, 1998
= Caudipteryx dongi Zhou and Wang, 2000
Late Barremian-Early Aptian, Early Cretaceous
Jianshangou member of Yixian Formation, Liaoning, China
Holotype- (NGMC 97-4-A) (890 mm) skull (76 mm), mandibles, cervical vertebae, cervical ribs, dorsal vertebrae, dorsal ribs, gastralia, twenty-two caudal vertebrae (first caudal 12 mm), chevrons, coracoid, sternal plate (36 mm), humeri (69 mm), radii, ulnae, scapholunare, semilunate carpal, distal carpal III, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, metacarpal III, ilia, pubes, ischia (77 mm), femora (147 mm), tibiae (188 mm), fibula, phalanx I-1, metatarsal II, phalanx II-1, phalanx II-2, partial pedal ungual II, metatarsal III (115 mm), fragmentary digit III, metatarsal IV, phalanx IV-1, contour feathers, remiges, retrices, gastroliths
Paratype- (NGMC 97-9-A) (725 mm) incomplete skull (79 mm), mandibles, hyoid, cervical vertebrae, cervical ribs, dorsal vertebrae, dorsal ribs, sacrum, caudal vertebrae, chevrons, scapulae (77 mm), coracoids, partial furcula, sternal plates, sternal ribs, humeri (70 mm), radii (57 mm), ulnae, semilunate carpal, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, metacarpal III, partial phalanx III-1, incomplete ilium, partial pubes, femora (149 mm), tibiae (182 mm), astragali, calcanea, metatarsal I, phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III (117 mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal V, contour feathers, retrices, remiges, gastroliths
Referred- (IVPP V11819) specimen including femur (149 mm) (Erickson et al., 2009)
(IVPP V 12344; holotype of Caudipteryx dongi) (896 mm) frontal, pterygoid, two cervical vertebrae, six dorsal vertebrae, dorsal ribs (100 mm), three uncinate processes (30 mm), gastralia, sacrum, eleven caudal vertebrae, chevrons, partial coracoid, sternal plates (25 mm), sternal ribs (35 mm), incomplete humeri (~73 mm), radii (~58 mm), ulnae (61 mm), semilunate carpal, scapholunare, ulnare, metacarpal I (13 mm), phalanx I-1 (25 mm), manual ungual I (15 mm), metacarpal II (29 mm), phalanx II-1 (18.5 mm), phalanx II-2 (25 mm), manual ungual II (18 mm), metacarpal III (27 mm), ilia (115 mm), pubes, ischia (73 mm), femora (146, 152 mm), tibiae (196 mm), fibula (181 mm), astragali, calcaneum, distal tarsal III, distal tarsal IV, metatarsal I (19 mm), phalanx I-1 (12 mm), pedal ungual I (11 mm), metatarsal II (112 mm), phalanx II-1 (25 mm), phalanx II-2 (16 mm), pedal ungual II (19 mm), metatarsal III (124 mm), phalanx III-1 (27 mm), phalanx III-2 (20 mm), phalanx III-3 (17 mm), pedal ungual III (20 mm), metatarsal IV (116 mm), phalanx IV-1 (15 mm), phalanx IV-2 (9 mm), phalanx IV-3 (7 mm), phalanx IV-4 (7 mm), pedal ungual IV (16 mm), metatarsal V (~36 mm), body feathers, remiges (182 mm), gastroliths (Zhou and Wang, 2000)
(LPM 0005) skull, mandibles, cervical series, cervical ribs, dorsal series, dorsal ribs, gastralia, sacral vertebrae, caudal series, chevrons, scapulae, coracoid, furcula, humeri, radii, ulnae, semilunate carpal, metacarpals I, phalanges I-1, manual unguals I, metacarpals II, phalanges II-1 (one partial), phalanx II-2, manual ungual II, metacarpals III, phalanges III-1, phalanges III-2, ilia, ischia, femora, tibiae, fibulae, astragali, calcaneum, metatarsals I, phalanges I-1, pedal ungual I, metatarsals II, phalanges II-1, phalanges II-2, pedal unguals II, metatarsals III, phalanges III-1, phalanges III-2, phalanges III-3, pedal unguals III, metatarsals IV, phalanges IV-1, phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal unguals IV, metatarsals V, remiges, retrices, body feathers, gastroliths (Feduccia and Czerkas, 2015)
(STM 4-3) specimen including dorsal vertebrae, dorsal ribs, scapula and coracoids (Zheng et al., 2014)
Diagnosis- (suggested) premaxilla sharply pointed anteriorly; quadratojugal posterior process slightly developed; manual ungual II larger than manual ungual I.
C. sp. nov. (Zhou, Wang, Zhang and Xu, 2000)
Late Valanginian-Middle Aptian, Early Cretaceous
Yixian Formation, Liaoning, China
Material- (BPM 0001) (852 mm) skull, mandibles, twelve cervical vertebrae, cervical ribs, nine dorsal vertebrae, dorsal ribs (~114 mm), uncinate processes, gastralia, sacrum, twenty-two caudal vertebrae, chevrons, scapulae (80 mm), coracoids (34 mm), sternal plates (~30 mm), sternal ribs (~38 mm), humeri (72 mm), radii (59 mm), ulnae (62 mm), semilunate carpal, scapholunare, ulnare, metacarpal I (11 mm), phalanx I-1 (25 mm), manual ungual I (16 mm), metacarpal II (28 mm), phalanx II-1 (17 mm), phalanx II-2 (24 mm), manual ungual II (15 mm), metacarpal III (25 mm), phalanx III-1, phalanx III-2, ilia (115 mm), pubes (~124 mm), ischia (72 mm), femora (145 mm), tibiae (188 mm), fibulae (188 mm), astragali, calcanea, distal tarsal III, distal tarsal IV, metatarsal I (16 mm), phalanx I-1 (13 mm), pedal ungual I (12 mm), metatarsal II (102 mm), phalanx II-1 (23 mm), phalanx II-2 (16 mm), pedal ungual II (19 mm), metatarsal III (113 mm), phalanx III-1 (24 mm), phalanx III-2 (19 mm), phalanx III-3 (15 mm), pedal ungual III (18 mm), metatarsal IV (107 mm), phalanx IV-1 (14 mm), phalanx IV-2 (8 mm), phalanx IV-3 (6 mm), phalanx IV-4 (4 mm), pedal ungual IV (14 mm), metatarsal V (30 mm), contour feathers, remiges, retrices, gastroliths
(IVPP V 12340) (836 mm) skull, mandibles, twelve cervical vertebrae, cervical ribs, nine dorsal vertebrae (d3 ~18 mm, d6 ~18 mm, d8 ~17 mm), dorsal ribs (120 mm), uncinate processes (26 mm), gastralia, sacrum, twenty-two caudal vertebrae, chevrons, scapulae (80 mm), coracoids (35 mm), sternal ribs (36 mm), humeri (69 mm), radii (~56 mm), ulnae (61 mm), semilunate carpal, scapholunare, ulnare, metacarpal I (11 mm), phalanx I-1 (26 mm), manual ungual I (16 mm), metacarpal II (28 mm), phalanx II-1 (19 mm), phalanx II-2 (24 mm), manual ungual II (15 mm), metacarpal III (23 mm), phalanx III-1, phalanx III-2, ilia (112 mm), pubes (~125 mm), ischia (~72 mm), femora (145 mm), tibiae (183 mm), fibulae (175 mm), astragali, calcanea, distal tarsal III, distal tarsal IV, metatarsal I (15 mm), phalanx I-1 (12 mm), pedal ungual I (12 mm), metatarsal II (102 mm), phalanx II-1 (22 mm), phalanx II-2 (14 mm), pedal ungual II (17 mm), metatarsal III (112 mm), phalanx III-1 (23 mm), phalanx III-2 (17 mm), phalanx III-3 (13 mm), pedal ungual III (18 mm), metatarsal IV (106 mm), phalanx IV-1 (12 mm), phalanx IV-2 (8 mm), phalanx IV-3 (6 mm), phalanx IV-4 (5 mm), pedal ungual IV, metatarsal V (31 mm), contour feathers, remiges, retrices, gastroliths
Diagnosis- (suggested) large premaxillary subnarial process; maxilla extended anteriorly with promaxillary fossa; external naris close to antorbital fenestra in length; jugal strongly concave posterodorsally; posterodorsal dentary process subequal in width to posteroventral process; posterodorsal dentary process longer than posteroventral process; no intramandibular joint; vomers do not extend past external nares?; ectopterygoid very thin and C-shaped; twelve cervical vertebrae?; ventral margin of coracoid irregular?; anterior margin of preacetabular process posteroventrally oriented.
Comparison of specimens- Five specimens of Caudipteryx have been described. NGMC 97-4-A, NGMC 97-9-A and BPM 0001 are referred to the type species, C. zoui (Ji et al., 1998; Zhou et al., 2000). IVPP V 12344 was referred to a new species, C. dongi (Zhou and Wang, 2000). IVPP V 12430 was referred simply to C. sp. (Zhou et al., 2000). Zhou and Wang differetiated C. dongi from C. zoui based on the smaller sternum and longer first metacarpal. Most differences I can see between the specimens are cranial, although this may be due to the fact the skulls are well illustrated, while the postcrania is not. Are these differences real or preservational? A large amount of the variety seems to be due to crushing and distortion. For instance, there is no way the lacrimal of IVPP V 12430 could have had such a small angle between its anterior and posterior processes in life. Similarily, the posterior postorbital process of BPM 0001 is much too long, as it would extend well past the quadrate when articulated. The anterior squamosal process of that specimen is much too large and bulbous, as it would reach through the postorbital and into the orbit. More evidence that distortion has occured might come from the asymmetry in specimens. The dorsal cranial elements (nasal, frontal, parietal) are often distorted and asymmetrical. The differences least likely to be due to distortion or individual variation support BPM 0001 and IVPP V 12430 being separate from NGMC 97-9-A. Characters these two specimens share not found in the latter are- premaxilla blunt anteriorly; large premaxillary subnarial process; maxilla extended anteriorly with promaxillary fossa; external naris close to antorbital fenestra in length; jugal strongly concave posterodorsally; quadratojugal posterior process not developed; posterodorsal dentary process subequal in width to posteroventral process; posterodorsal dentary process longer than posteroventral process; no intramandibular joint. Most of the postcranium is not figured in sufficient detail to determine morphological differences in specimens. The differing number of reported cervical vertebrae might be due to misinterpretation, as Zhou et al. state "there are estimated twelve cervical vertebrae". The coracoid has a smoothly rounded ventral border in NGMC 97-9-A, unlike the irregular border of BPM 0001, although the significance of this is uncertain. The orientation of the anterior preacetabular edge differs in IVPP V 12344 and IVPP V 12430, but as the skull of the former is fragmentary, it cannot be determined if this is correlated with the cranial differences noted above. Contra Zhou and Wang, no significant differences in postcranial ratios is evident. Most ratios vary within a few percentage points of each other, so fall within the expected range of individual variation. The sternal plates are 24% of femoral length in the holotype of C. zoui, 17% in the holotype of C. dongi and an intermediate 21% in BPM 0001. A three percent difference in size does not seem to fall outside the range of individual variation. The first metacarpal of BPM 0001 and IVPP V 12430 is 39% of metacarpal II length. In IVPP V 12344, the ratio is 45%. This difference might be considered diagnostic if not for NGMC 97-4-A, which has a 42% ratio. Although stated to be "about .4" in Zhou and Wang, this figure comes from Ji et al., who only measured to the tenths place. The exact ratio, as mentioned above, is intermediate between the more divergent specimens. Once again, the 3% difference is considered insufficient to diagnose a species. The ilium is much shorter in NGMC 97-4-A (69% of femoral length) than in BPM 0001, IVPP V 12344 and IVPP V 12430 (77-79%). This is due to the broken anterior edge in the former specimen however, as can be seen in the specimen (pers. obs.). The only potentially significant proportional difference between specimens is- manual ungual I vs. manual ungual II (106% in BPM 0001 and IVPP V 12430, 84% in IVPP 12344). Although the first manual ungual of NGMC 97-4-A is incomplete, it was much smaller than manual ungual II, so seems to match IVPP V 12344 better. It therefore seems that BPM 0001 and IVPP V 12430 share several cranial characters not seen in NGMC 97-9-A; NGMC 97-9-A has a slightly different coracoid morphology than BPM 0001; BPM 0001 and IVPP V 12430 have a different preacetabular morphology than IVPP V 12344; and that BPM 0001 and IVPP V 12430 have different manual ungual ratios than IVPP V 12344 and NGMC 97-4-A. Two groups of specimens are suggested by these differences- IVPP V 12344, NGMC 97-4-A and NGMC 97-9-A are one group, while BPM 0001 and IVPP V 12430 are in the other. The inclusion of IVPP V 12344 and NGMC 97-4-A with NGMC 97-9-A is far from certain, but the fact they all differ from the other two specimens and that the latter two have similar ungual ratios suggests this may be the case. Are these differences due to ontogenetic, sexual or taxonomic variation? The minute size variation (femora vary between 145-152 mm) suggests it is not ontogenetic. Settling whether two sexes or species are involved is not easily resolved with only five specimens to work with, all from different localities. One potential way to decide this would be if the groups are not sister groups in a phylogenetic analysis.
Phylogenetic relationships- Sereno (1999) was the first to recover Caudipteryx as an oviraptorosaur which has been the consensus since 2002, but a few other possibilities have been proposed over the years.
Caudipteryx in Avialae? Ji et al. (1998) first included Caudipteryx in Chiappe's bird matrix, also including alvarezsaurids and Protarchaeopteryx, with Velociraptor as an outgroup. Note this doesn't allow Caudipteryx to fall outside of Eumaniraptora. Also note it was only examined with characters that were thought to be useful for analyzing Avialae. With these caveats in mind, Ji et al. found it to be an avialan based on two characters- unserrated teeth; dorsal premaxillary process reaches to anterior border of antorbital fossa. As only the premaxilla is toothed in Caudipteryx, and premaxillary teeth are plesiomorphically unserrated in maniraptoriforms (only derived dromaeosaurids and derived troodontids have serrations; Protarchaeopteryx was misscored as having them), this doesn't support placing Caudipteryx in Avialae instead of Oviraptorosauria. The second character is also seen in caenagnathoids (Anzu, Avimimus, oviraptorids), though absent in Incisivosaurus.
Those who doubted the dinosaur-bird link have always said Caudipteryx is a bird due to its unambiguous remiges and retrices. Originally, this meant separating it from other non-avialan maniraptoriforms, which they viewed as dinosaurs. Since 2002 however, as more maniraptoriforms are discovered with remiges and retrices, workers such as Feduccia and Martin have allowed oviraptorosaurs, dromaeosaurids, troodontids, and possibly even alvarezsaurids and ornithomimosaurs to be birds as well (though they still insist therizinosaurs are sauropodomorphs). Thus their arguments for placing Caudipteryx as a bird (e.g. Martin and Czerkas, 2000; Geist and Feduccia, 2000; Ruben and Jones, 2000) are no longer valid, as they now think some taxa which lack these bird-like characters (e.g. Velociraptor) are birds anyway. Similarily, Feduccia et al. (2005) and Martin (2004) now agree Caudipteryx is a basal oviraptorosaur, though their placement of oviraptorosaurs and other maniraptorans outside of Theropoda remains incorrect.
Martin and Czerkas (2000) argued Caudipteryx was a sauriurine more closely related to Confuciusornis than to Archaeopteryx. It is uncertain whether they considered enantiornithines to be the sister taxon to Confuciusornis or to Caudipteryx+Confuciusornis. Of their supporting characters, a reduced fibula, reduced calcaneum, and "evidence of pygostyle formation" are not present in Caudipteryx. A reduced hypopubic cup is a fictional character, as no bird has a hypopubic cup. The external mandibular fenestra and ball-shaped femoral head are plesiomorphies only absent in a few basal paravians like Archaeopteryx, but present in at least some enantiornithines in addition to confuciusornithids. On the other hand, the enlarged premaxilla, reduced maxilla, toothless maxilla and dentary and shortened tail are indeed shared with Confuciusornis but not Archaeopteryx, but are also found in caenagnathoids. Martin (2004) even suggested Caudipteryx and other oviraptorosaurs may be most closely related to confuciusornithids. This is highly unparsimonious though, as oviraptorosaurs lack numerous paravian/eumaniraptoran, avialan, avebrevicaudan and pygostylian characters found in confuciusornithids; basal oviraptorosaurs such as Incisivosaurus and Caudipteryx lack most of the confuciuornithid-like characters found in derived oviraptorosaurs; and confuciusornithids in turn lack most oviraptoriform and oviraptorosaur characters.
Lu et al. (2002) found Caudipteryx to be closer to birds than Archaeopteryx but basal to paraphyletic alvarezsaurids, oviraptorids and ornithothoracines. Similar to Ji et al.'s analysis, this was based on Chiappe's bird matrix so has the same issues, except this time almost all alvarezsaurid characters were excluded and Oviraptorosauria was added as an OTU though without many characters linking Caudipteryx to it. So this matrix again forces Caudipteryx, alvarezsaurids and now oviraptorosaurs to be avialans, which is unparsimonious when non-paravian outgroups are used.
Caudipteryx in Paraves? Xu et al. (1999) resolved Caudipteryx as a paravian in a trichotomy with Protarchaeopteryx and Troodontidae+Eumaniraptora. Xu et al. (2000) used 86 of the same characters plus three new ones, added Microraptor and removed Tyrannosauridae and Unenlagia. Their tree was slightly more resolved, as Caudipteryx was found to be outside a clade consisting of Protarchaeopteryx and Troodontidae+Eumaniraptora. However, both studies supported the paravian status of Caudipteryx using the same ten characters. Of these, a deep jugal, unfused interdental plates, less than 11 caudal vertebrae with transverse processes, metacarpal I less than 33% of metacarpal II in length, pubic foot projects posteriorly only and posterior trochanter present are misscored in Caudipteryx. Oviraptorids and troodontids are miscoded as lacking a proximodistally elongate coracoid, which is actually a pennaraptoran synapomorphy. Most deinonychosaurs (except Archaeopteryx, Ningyuanosaurus, Eosinopteryx and Jinfengopteryx) and jeholornithids have more than 25 caudal vertebrae, so the low number in Caudipteryx is not a paravian character (and is shared with e.g. Similicaudipteryx). A radius less than 70% of ulnar width is shared with e.g. Protarchaeopteryx, Similicaudipteryx and Anzu and is highly homoplasious in basal Paraves. However, mid and distal chevrons dorsoventrally flattened (defined as in Hartman et al. 2019 as 'Chevrons - anteroposterior length >75% of dorsoventral depth by chevron 10') is shared with almost all paravians to the exclusion of oviraptorosaurs.
Caudipteryx outside Pennaraptora? Holtz (2001) found Caudipteryx to either be sister to a therizinosaur plus oviraptorosaur clade or a basal pennaraptoran. Gishlick (2002) found Caudipteryx to be in a trichotomy with therizinosaurs and Pennaraptora, but the matrix only contained forelimb characters.
References- Ji, Currie, Norell and Ji, 1998. Two feathered dinosaurs from northeastern China. Nature. 393, 753-761.
Sereno, 1999. The evolution of dinosaurs. Science. 284, 2137-2147.
Xu, Wang and Wu, 1999. A dromaeosaurid dinosaur with filamentous integument from the Yixian Formation of China. Nature. 401, 262-266.
Jones, Farlow, Ruben, Henderson and Hillenius, 2000. Cursoriality in bipedal archosaurs. Nature. 406, 716-718.
Geist and Feduccia, 2000. Gravity defying behaviors: Identifying models for protoaves. American Zoologist. 40(4), 664-675.
Martin and Czerkas, 2000. The fossil record of feather evolution in the Mesozoic. American Zoologist. 40(4), 687-694.
Ruben and Jones, 2000. Selective factors associated with the origin of fur and feathers. American Zoologist. 40(4), 585-596.
Xu, Zhou and Wang, 2000. The smallest known non-avian theropod dinosaur. Nature. 408, 705-708.
Zhou and Wang, 2000. A new species of Caudipteryx from the Yixian Formation of Liaoning, northeast China. Vertebrata PalAsiatica. 38(2), 113-130.
Zhou, Wang, Zhang and Xu, 2000. Important features of Caudipteryx - evidence from two nearly complete new specimens. Vertebrata PalAsiatica. 38(4), 241-254.
Holtz, 2001. Arctometatarsalia revisited: The problem of homplasy in reconstructing theropod phylogeny. In Gauthier and Gall (eds.). New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom. Yale University Press. 99-122.
Ruben and Jones, 2001. Feathered dinosaurs and other myths: a cold, hard look at reality. Journal of Morphology. 248(3), 278.
Christiansen and Bonde, 2002. Limb proportions and avian terrestrial locomotion. Journal of Ornithology. 143, 356-371.
Gishlick, 2002. The functional morphology of the forelimb of Deinonychus antirrhopus and its importance for the origin of avian flight. Unpublished PhD thesis. Yale University, 142 pp.
Lu, Dong, Azuma, Barsbold and Tomida, 2002. Oviraptorosaurs compared to birds. In Zhou and Zhang (eds.). Proceedings of'the 5th Symposium of the Society of Avian Paleontology and Evolution. 175-189.
Martin, 2004. A basal archosaurian origin for birds. Acta Zoologica Sinica. 50(6), 978-990.
Dyke and Norell, 2005. Caudipteryx as a non-avialan theropod rather than a flightless bird. Acta Palaeontologica Polonica. 50(1), 101-116.
Feduccia, Lingham-Soliar and Hinchliffe, 2005. Do feathered dinosaurs exist? Testing the hypothesis on neontological and paleontological evidence. Journal of Morphology. 266(2), 125-166.
Erickson, Rauhut, Zhou, Turner, Inouye, Hu and Norell, 2009. Was dinosaurian physiology inherited by birds? Reconciling slow growth in Archaeopteryx. PLoS ONE. 4(10), e7390.
Zheng, O'Connor, Wang, Wang, Zhang and Zhou, 2014. On the absence of sternal elements in Anchiornis (Paraves) and Sapeornis (Aves) and the complex early evolution of the avian sternum. Proceedings of the National Academy of Sciences. 111(38), 13900-13905.
Feduccia and Czerkas, 2015. Testing the neoflightless hypothesis: Propatagium reveals flying ancestry of oviraptorosaurs. Journal of Ornithology. 156(4), 1067-1074.

"Xingtianosaurus" Qiu, Wang, Wang, Li, Zhang and Ma, 2019
"X. ganqi" Qiu, Wang, Wang, Li, Zhang and Ma, 2019
Early Aptian, Early Cretaceous
Dawangzhangzi Beds of Yixian Formation, Liaoning, China
Material- (IVPP V13390) four dorsal centra, dorsal rib fragments, (?)uncinate processes, gastralia, anterior sacral centrum, about eighteen caudal vertebrae and centra, nine chevrons, partial scapulae (~56.2 mm), (?)sternal plate (~32.7 mm), humeri (one incomplete, one partial; 72.4 mm), radii (69.9, 63.3 mm), ulnae (one incomplete; 73.5, 69.5 mm), scapholunare, (?)ulnare, semilunate carpals, metacarpals I (16.6, 15.4 mm), phalanges I-1 (one partial; 25.5 mm), manual ungual I (impression), metacarpals II (one incomplete; 40.5 mm), proximal phalanx II-1, distal phalanx II-2, partial manual unguals II (21.2, ~19.7 mm), metacarpals III (one incomplete; 37.2 mm), proximal phalanx III-1, distal phalanx III-3, manual unguals III (15.4, 15.6 mm), partial ilium (~89.8 mm), pubes (~107 mm), ischium (~54.6 mm), femora (127, 120.4 mm), incomplete tibiae (171.3 mm), incomplete fibulae, astragalus, two distal tarsals, metatarsals I (9.8, 10.2 mm), phalanx I-1 (19 mm), pedal ungual I (14.6 mm), metatarsals II (80.4, 78.5 mm), phalanges II-1 (28.3, 27.8 mm), phalanges II-2 (22.6, 23.8 mm), pedal unguals II, metatarsals III (89.7, 90.2 mm), phalanges III-1 (27, 27 mm), phalanges III-2 (20.9, 20.7 mm), phalanges III-3 (20.3, 21.4 mm), pedal unguals III (17.2, 15 mm), metatarsals IV (82.8, 82.9 mm), phalanges IV-1 (17.3, 17.6 mm), phalanges IV-2 (14.5, 14.3 mm), phalanges IV-3 (12.7, 12.8 mm), phalanges IV-4 (16, 14.7 mm), pedal unguals IV (16, 15.8 mm), metatarsal V (25.6 mm), remiges
Other diagnoses- Of the listed characters in Qiu et al.'s (2019) diagnosis, none definitely distinguish this taxon from Protarchaeopteryx.
- "small pleurocoel close to the dorsal edge of the lateral surface of the dorsal vertebral centrum", pleurocoels are also present in Protarchaeopteryx's posterior dorsals (unlike Caudipteryx) although their size and position are unreported.
- "humerus longer than the scapula" is unknown in Protarchaeopteryx as only the proximal scapula is preserved.
- "proportionally long ulna (as long as humerus)" with an ulnohumeral ratio of 101%, which is larger than ~86% in Protarchaeopteryx. This is actually due to the longer humerus in Protarchaeopteryx, as the femur and ulna are about the same length in each specimen. One interesting possibility is that previous authors were mistaken in identifying the proximal humerus of Protarchaeopteryx, as the purported fragment of is indistinct and even has a hole which could be a coracoid foramen.
- "relatively small radiale [= scapholunare] angle (39°, compared to >48° in other oviraptorosaurs with known radiale angle)" compared to 26° in Protarchaeopteryx.
- "extremely short metacarpal I (<40% length of the metacarpal II)" is actually 41%, compared to ~39% in Protarchaeopteryx.
- "small ligament pits on the manual phalanges" doesn't seem distinct from some Caudipteryx (NGMC 97-9-A) in the one phalanx illustrated as preserving them (II-2), but does seem larger in at least I-1 and III-3 of Protarchaeopteryx. Qiu et al.'s Caudipteryx example (their Fig. 7c) is IVPP V12430, which does have a large pit on II-2.
Comments- This was described by Qiu et al. on April 25 2019 as a new taxon of caudipterid. However, this paper has no mention of ZooBank and as of February 6 2020 "Xingtianosaurus" lacks an entry on the ZooBank website. Thus according to ICZN Article 8.5.3 (an electronic work must "be registered in the Official Register of Zoological Nomenclature (ZooBank) (see Article 78.2.4) and contain evidence in the work itself that such registration has occurred"), "Xingtianosaurus ganqi" Qiu et al., 2019 is a nomen nudum that will only be technically valid pending action on behalf of the authors or ICZN as its journal is not published physically.
If Protarchaeopteryx's humerus has been correctly interpreted, it's much longer than "Xingtianosaurus"', but the altter's greater scapholunare angle and smaller manual ligament pits on at least two phalanges may be outside of individual variation. Yet the sternal plates would certainly seem to be very different. Protarchaeopteryx's are a rounded rectangle, while "Xingtianosaurus"' are illustrated as being drop-shaped with a notch that creates anterolateral and posterolateral processes. Yet the medial edge is along a crack in the matrix, with similar texture to the sternal bone on the other side in some areas, and the entire anterior edge is hidden by ribs. Only a short section representing about a fourth of the supposed lateral notch is exposed, and this could be breakage. Indeed, with only a short curved portion of the edge obviously visible, the whole bone could easily be one of the otherwise missing coracoids. Thus definitely distinguishing the taxa may involve the proper identification of elements beyond what published photos allow.
Qiu et al. (2019) used a version of Maryanska's oviraptorosaur matrix and recovered "Xingtianosaurus" as a caudipterid. Adding it to Hartman et al.'s analysis also recovers it as a caudipterid.
Reference- Qiu, Wang, Wang, Li, Zhang and Ma, 2019. A new caudipterid from the Lower Cretaceous of China with information on the evolution of the manus of Oviraptorosauria. Scientific Reports. 9:6431. DOI: 10.1038/s41598-019-42547-6

Incisivosaurus Xu, Cheng, Wang and Chang, 2002
I. gauthieri Xu, Cheng, Wang and Chang, 2002
= Protarchaeopteryx gauthieri (Xu, Cheng, Wang and Chang, 2002) Senter, Barsbold, Britt and Burnham, 2004
Late Valanginian-Hauterivian, Early Cretaceous
Lujiatun Beds of Yixian Formation, Liaoning, China
Holotype- (IVPP V13326) skull (100 mm), incomplete mandibles, partial cervical vertebra
Diagnosis- (after Xu et al., 2002) large high-angled wear facets on the mesial margins of the teeth; contact between the accessory ventral flanges of the pterygoids.
(after Senter et al., 2004) nine maxillary teeth; eight or nine dentary teeth.
Other diagnoses- Xu et al. (2002) also included the following characters in their diagnosis- large incisciform first premaxillary tooth (also in Protarchaeopteryx); much smaller, subconical second to fourth premaxillary teeth (plesiomorphic for Oviraptorosauria); very small lanceolate maxillary teeth (plesiomorphic for Oviraptorosauria); triradiate palatine with very short maxillary process (plesiomorphic for Oviraptoriformes); longitudinal crest on the ventral surface of the basisphenoid (plesiomorphic for coelurosaurs). The "subsidiary ectopterygoid fenestra" they note seems to be merely a reduced subsidiary palatal fenestra shifted between the ectopterygoid and palatine as in oviraptorids.
Comments- Balanoff et al. (2009) note the posterior mandibular fragment is lost.
References- Xu, Cheng, Wang and Chang, 2002. An unusual oviraptorosaurian dinosaur from China. Nature. 419, 291-293.
Senter, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae. Bulletin of Gunma Museum of Natural History. 8, 1-20.
Balanoff, Xu, Matsufune, Kobayashi and Norell, 2007. Endocranial anatomy of a primitive oviraptorosaur, Incisivosaurus gauthieri, (Theropoda: Dinosauria). Journal of Vertebrate Paleontology. 27(3), 43A.
Balanoff, Xu, Kobayashi, Matsufune and Norell, 2009 online. Incisivosaurus gauthieri, Digital Morphology. http://digimorph.org/specimens/Incisivosaurus_gauthieri/
Balanoff, Xu, Kobayashi, Matsufune and Norell, 2009. Cranial osteology of the theropod dinosaur Incisivosaurus gauthieri (Theropoda: Oviraptorosauria). American Museum Novitates. 3651, 35 pp.
Balanoff, 2011. Oviraptorosauria: Morphology, phylogeny, and endocranial evolution. PhD thesis. Columbia University. 522 pp.

 Yuanyanglong Hao, Li, Wang, Wang, Ma, Qinggele, King, Pei, Zhao and Xu, 2024 online 
Y. bainian Hao, Li, Wang, Wang, Ma, Qinggele, King, Pei, Zhao and Xu, 2024 online
Etymology- "Genus name from 'Yuanyang', mandarian for lovebirds (Aix galericulata), which symbolizes living in pairs forever, referring to the paired individuals in one block; and 'long', a transliteration of Chinese word for dragon. The species name 'bainian' means 'a hundred years' in Chinese, in commemoration of exactly 100 years from the initial erection of the original oviraptorosaur species (Chirostenotes pergracilis Gilmore 1924; Oviraptor philoceratops Osborn 1924)."
Aptian-Albian, Early Cretaceous
Maortu, Miaogou Formation, Inner Mongolia, China 
Holotype- (PV02476-1; field number MET-21-037) (11.80 kg based on Campione et al., 2014) two anterior middle dorsal centra, six posterior dorsal neural spines, eight right dorsal ribs, two right ?unicinate process fragments, two anterior synsacral centra, fifth and sixth sacral neural spines, first and second caudal vertebrae, second chevron, right scapula, incomplete right coracoid, right sternum?, fragmentary sternal ribs, right humerus, proximal right radius, proximal right ulna, right ilium, right pubis, distal left pubis, ischia, femora, left tibiotarsus, left proximal fibula, distal tarsal II fused to proximal metatarsal II, proximal metatarsal III, distal tarsal IV fused to proximal metatarsal IV, gastroliths
Paratype- (PV02476-2; field number MET-21-037) partial skull, three posteriormost dorsal vertebrae, first-fifth synsacral vertebrae, partial pelvis, left femur, right tibiotarsus, right distal tarsal II fused to proximal metatarsal II, proximal right metatarsal III, right distal tarsal IV fused to proximal metatarsal IV
Diagnosis- (after Hao et al., 2024 online; autapomorphies only) preacetabular process of ilium approximately 40% longer than postacetabular process; hook-like anteroventral process of ilium extending to level of dorsal acetabular margin; reduced anterior pubic boot shorter than hooked posterior boot; slender tibiotarsus (distal end transverse width to whole bone length ratio about 0.08).
Comments- Discovered in summer 2021. Hao et al. (2024 online) state "The well-fused sacrals, tibiotarsus and tarsometatarsus in two individuals are generally considered evidence of adulthood", thus they "tentatively consider PV 02476 to be representative of the adult body size of Yuanyanglong." Gastralia are also preserved.
Hao et al. (2024 online) used Funston's oviraptorosaur matrix to recover Yuanyanglong sister to Avimimidae + Caenagnathoidea.
References- Hao, Li, Wang, Wang, Ma, Qinggele, King, Pei, Zhao and Xu, 2024 online. A new oviraptorosaur from the Lower Cretaceous Miaogou Formation of western Inner Mongolia, China. Cretaceous Research. DOI: 10.1016/j.cretres.2024.106023

Caenagnathoidea Sternberg, 1940 sensu Sereno, 1999b
Definition- (Caenagnathus collinsi + Oviraptor philoceratops) (Maryanska, Osmólska and Wolsan, 2002; modified from Sereno, 1999b)
Other definitions- (Chirostenotes pergracilis + Oviraptor philoceratops) (Sereno, online 2005)
(Caenagnathus collinsi + Oviraptor philoceratops + Avimimus portentosus) (Hendrickx, Mateus, Araújo and Choiniere, 2019)
= Oviraptorosauria sensu Padian, Hutchinson and Holtz, 1999
Definition- (Oviraptor philoceratops + Chirostenotes pergracilis) (modified)
= Oviraptoroidea Barsbold, 1976 sensu Sereno, 1999a
Definition- (Oviraptor philoceratops + Caenagnathus collinsi) (modified)
= Caenagnathoidea sensu Sereno, online 2005
Definition- (Chirostenotes pergracilis + Oviraptor philoceratops)
= Caenagnathoidea sensu Hendrickx, Mateus, Araújo and Choiniere, 2019
Definition- (Caenagnathus collinsi + Oviraptor philoceratops + Avimimus portentosus)
= Edentoraptora Funston, Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020
Comments- This node covered known oviraptorosaurs until Sereno (1999a) identified Caudipteryx as a basal oviraptorosaurian and proposed Oviraptoroidea as a name. However, according to ICZN rules Caenagnathoidea was implicitly named in 1940 when Sternberg erected Caenagnathidae, so it must have priority over Oviraptoroidea which was implicitly created in 1976 by Barsbold. Thus (though without explanation in the text) Sereno (1999b) next used Caenagnathoidea for this node, with its eponymous genera implied by the families it contains in the cladogram.
Funston et al. (2020) noted "a well-supported (decay index 2) group of Avimimus and Caenagnathoidea, referred to here as Edentoraptora based on the ubiquitous absence of teeth in these animals." No phylogenetic definition was provided and the group has the same content as Caenagnathoidea in this site's topology based on Hartman et al. where Avimimus is a caenagnathid. Forcing Avimimus outside Caenagnathoidea in that matrix and thus making Edentoraptora a unique clade takes ten additional steps.
Caenagnathoidea defined- Maryanska et al. (2002) specified the defining genera and species, but Sereno (online 2005) tried to replace Caenagnathus with Chirostenotes. This is a poor decision, as the taxa are not definitely synonymous. Chirostenotes pergracilis and Elmisaurus elegans co-occur in the same formation, and the only reason Caenagnathus is synonymized with pergracilis instead of elegans is size. Until taxonomic problems are solved for caenagnathids, it's best to associate the family with its eponymous species. Hendrickx et al. (2019) added Avimimus to the definition as an internal specifier, but this is unneccesary and indeed whether Avimimus is a caenagnathoid is an active problem in oviraptorosaur phylogeny.
References- Sternberg, 1940. A toothless bird from the Cretaceous of Alberta. Journal of Paleontology. 14(1), 81-85.
Barsbold, 1976. On a new Late Cretaceous family of small theropods (Oviraptoridae fam. n.) of Mongolia. Doklady Akademia Nauk SSSR. 226, 685-688.
Padian, Hutchinson and Holtz, 1999. Phylogenetic definitions and nomenclature of the major taxonomic categories of the carnivorous Dinosauria (Theropoda). Journal of Vertebrate Paleontology. 19(1), 69-80.
Sereno, 1999a. The evolution of dinosaurs. Science. 284, 2137-2147.
Sereno, 1999b. A rationale for dinosaurian taxonomy. Journal of Vertebrate Paleontology. 19(4), 788-790.
Maryanska, Osmólska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Sereno, online 2005. Stem Archosauria - TaxonSearch. http://www.taxonsearch.org/dev/file_home.php [version 1.0, 2005 November 7]
Hendrickx, Mateus, Araújo and Choiniere, 2019. The distribution of dental features in non-avian theropod dinosaurs: Taxonomic potential, degree of homoplasy, and major evolutionary trends. Palaeontologia Electronica. 22.3.74, 1-110.
Funston, Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020. A new two-fingered dinosaur sheds light on the radiation of Oviraptorosauria. Royal Society Open Science. 7: 201184.

undescribed Caenagnathoidea (Buckley, 2002)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US
Material- (NS.1563.018) two partial manual unguals, partial tibia, proximal fibula, partial astragalus, (metatarsal II ~390 mm) distal metatarsal III, partial metatarsal shaft, incomplete pedal phalanx III-1, partial pedal phalanx (Buckley, 2002)
(NS.32001.077) metatarsal III, metatarsal IV (Buckley, 2002)
manual ungual (Holtz, Williams, Tremaine and Matthews, 2014)
Comments- The NS specimens were identified as Elmisaurus in the abstract, but as caenagnathid or oviraptorid in the poster. They may belong to Anzu or Elmisaurus? sp. both known from the formation.
References- Buckley, 2002. New material of Elmisaurus (Theropoda, Elmisauridae) from the Late Cretaceous Hell Creek Formation of southeastern Montana. Journal of Vertebrate Paleontology. 22(3), 39A.
Holtz, Williams, Tremaine and Matthews, 2014. New additions to the Hell Creek Formation (Upper Maastrichtian) vertebrate fauna of Carter County, Montana. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 149.

undescribed caenagnathoid (Suzuki and Watabe, 2000)
Cenomanian-Turonian, Late Cretaceous
Bayn Shire, Baynshiren Formation, Mongolia
Material- (IGM coll.; 960805 BS WTB/M9; two hatchling individuals) articular-surangular-coronoid, two eggs, eggshell fragments
Comments- This was discovered in August 4-5 1995 and announced by Suzuki and Watabe (2000) as "A large-sized egg-nest was discovered from the lower mudstone layer. Eggs in the nest have elongated shape with prominent and linear surface ornamentation, with thick shell. When we screened the eroded sediments on the surface of surrounding area of the nest, we could find many embryonic bones together with shell debris of eggs from the nest." Watabe and Suzuki (2000) reported that the site was visited the next year between July 23 and August 8, and "many embryonic
bones were picked up together with a huge amount of egg debris. The eggshell was of oogenus Macroolithus with prominent ornaments, belonging to oofamily Elongatoolithidae. Iijima et al. (2011) reported in an abstract that the "transversely wide and convex articular facet, which is elevated above the remaining ramus, for the quadrate in lateral view" is a ceanagnathoid character.
References- Suzuki and Watabe, 2000. Report on the Japan - Mongolia Joint Paleontological Expedition to the Gobi desert, 1995. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 45-57.
Watabe and Suzuki, 2000. Report on the Japan - Mongolia Joint Paleontological Expedition to the Gobi desert, 1996. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 58-68. 60
Iijima, Sato, Watabe, Tsogtbaatar and Ariunchimeg, 2011. Bone bed of baby oviraptorosaur and hadrosauroid dinosaurs from the Bayanshiree Formation (Late Cretaceous) in southeastern Mongolia. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 130.

Caenagnathidae Sternberg, 1940
Definition- (Caenagnathus collinsi <- Oviraptor philoceratops) (Maryanska, Osmólska and Wolsan, 2002; modified from Sereno, 1998)
Other definitions- (Chirostenotes pergracilis + Chirostenotes elegans + Elmisaurus rarus + Caenagnathasia martinsoni + BHM 2033) (Sues, 1997)
(Chirostenotes pergracilis <- Oviraptor philoceratops) (Sereno, online 2005; modified from Padian, Hutchinson and Holtz, 1999)
= "Elmisauridae" Osmólska, 1980
= Elmisauridae Osmólska, 1981
= Kuszholiidae Nessov, 1992
= Caenagnathidae sensu Padian et al., 1999
Definition- (Chirostenotes pergracilis <- Oviraptor philoceratops) (modified)
Comments- Elmisaurids were first listed in Osmólska's (1980) Table III as carnivores of the Nemegt Formation represented by two individuals of one taxon, but as Elmisaurus itself was still listed as "small theropod gen. et sp. nov.", the name was unofficial. It is later listed as "a new family [Osmólska, in press]" with "Osmólska H. (in press). - Coossified tarsometatarsi in theropod dinosaurs and their bearing on the problem of bird origin." in the bibliography.
Ex-caenagnathids- Possible caenagnathid material from the Yalovach Formation of Tadjikistan (Ryan, 1997) is probably based on reports of oviraptorids by Nessov (1995), which probably belongs to therizinosaurs instead (Alifanov and Averianov, 2006). Reported caenagnathid forelimb elements from the Densus-Ciula Formation of Romania (Csiki and Grigorescu, 2005) have since been referred to the paravian Balaur (Csiki et al., 2010).
References- Sternberg, 1940. A toothless bird from the Cretaceous of Alberta. Journal of Paleontology. 14(1), 81-85.
Osmólska, 1980. The Late Cretaceous vertebrate assemblages of the Gobi Desert, Mongolia. Memoires de la Societe Geologique de France. 139, 145-150.
Osmólska, 1981. Coossified tarsometatarsi in theropod dinosaurs and their bearing on the problem of bird origins. Palaeontologia Polonica. 42, 79-95.
Nessov, 1992. Review of localities and remains of Mesozoic and Paleogene birds of the USSR and the description of new findings. Russkii Ornitologicheskii Zhurnal. 1(1), 7-50.
Nessov, 1995. Dinosaurs of northern Eurasia: New data about assemblages, ecology and paleobiogeography. Scientific Research Institute of the Earth's Crust, St. Petersburg State University, St. Petersburg, Russia. 156 pp.
Ryan, 1997. Middle Asian Dinosaurs. In Currie and Padian (eds.). Encyclopedia of Dinosaurs. Academic Press. 442-444.
Sues, 1997. On Chirostenotes, a Late Cretaceous oviraptorosaur (Dinosauria: Theropoda) from western North America. Journal of Vertebrate Paleontology. 17(4), 698-716.
Sereno, 1998. A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen. 210(1), 41-83.
Padian, Hutchinson and Holtz, 1999. Phylogenetic definitions and nomenclature of the major taxonomic categories of the carnivorous Dinosauria (Theropoda). Journal of Vertebrate Paleontology. 19(1), 69-80.
Maryanska, Osmólska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Csiki and Grigorescu, 2005. A new theropod from Tustea: Are there oviraptorosaurs in the Upper Cretaceous of Europe? Kaupia. 14, 78.
Sereno, online 2005. Stem Archosauria - TaxonSearch. http://www.taxonsearch.org/dev/file_home.php [version 1.0, 2005 November 7]
Alifanov and Averianov, 2006. On the finding of ornithomimid dinosaurs (Saurischia, Ornithomimosauria) in the Upper Cretaceous beds of Tajikistan. Paleontological Journal. 40(1), 103-108.
Csiki, Vremir, Brusatte and Norell, 2010. An aberrant island-dwelling theropod dinosaur from the Late Cretaceous of Romania. Proceedings of the National Academy of Sciences. 107(35), 15357-15361.

Luoyanggia Lu, Xu, Jiang, Jia, Li, Yuan, Zhang and Ji, 2009
L. liudianensis Lu, Xu, Jiang, Jia, Li, Yuan, Zhang and Ji, 2009
Aptian-Albian, Early Cretaceous
Haoling Formation, Henan, China
Holotype- (41HIII-00010) dentaries
....(41HIII-00011; field number KLR 07-62-44a) incomplete ilium, ischium
....(field number KLR 07-62-28a-16) metatarsal II (175), metatarsal III (190 mm), metatarsal IV (177 mm)
....(field number KLR 07-62-49-1) ilium, incomplete pubis (160 mm), ischium (100 mm)
Other diagnoses- Lu et al. (2009) listed three characters in their diagnosis, but all are plesiomorphic- dentary not decurved; dentary symphysis V-shaped; ischium only slightly concave dorsally.
Comments- While originally referred to the Mangchuan Formation, this was redefined to include three formations by Xu et al. in 2012.
While Lu et al. (2009) referred Luoyanggia to Oviraptoridae, they seemingly mean Oviraptorosauria as they refer Chirostenotes to that family and state "Luoyanggia may have a certain relationship with Caudiptery"x. Funston and Currie (2016) added it to Maryanska's oviraptorosaur analysis and found it resolve sister to Similicaudipteryx as the most basal oviraptorosaurs (note their published results did not find their shortest trees). Hartman et al. (2019) recovered it as a caenagnathine caenagnathid sister to Anzu, but adding more taxa places it less specifically as a caenagnathid outside the Elmisaurus+Avimimus clade.
Reference- Lu, Xu, Jiang, Jia, Li, Yuan, Zhang and Ji, 2009. A preliminary report on the new dinosaurian fauna from the Cretaceous of the Ruyang Basin, Henan province of Central China. Journal of the Paleontological Society of Korea. 25(1), 43-56.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

undescribed caenagnathid (Britt, Chure, Currie, Holmes, Theurer and Scheetz, 2021)
Late Albian, Early Cretaceous
DNM-16, Mussentuchit Member, Cedar Mountain Formation, Uhat, US
Material- (MCZ 3040) (~3 m, 4 year old subadult) posterior dorsal vertebra, rib fragments, incomplete femur, tibia, astragalocalcaneum, couple partial metatarsals, phalanx II-1, pedal phalanges
Comments- Found in the mid 1970s, this was presented by Britt et al. at SVP 2021. They wrote "1) camellate pneumatization of posterior dorsal vertebrae, 2) extremely high, parallel sided astragalar ascending process, 3) diminutive calcaneum fused to the astragalus, and 4) adaptations for extreme retraction and flexion in pedal Digit II" ..." suggests affinities with the Troodontidae." However, the characters of pedal phalanx II-1 are also present in Elmisaurus, the other characters are also present in oviraptorosaurs, and e.g. the pleurocoelous posterior dorsal is unknown in troodontids but consistant with caenagnathoids. Inclusion in Hartman et al.'s maniraptoromorph matrix results in a placement within Caenagnathidae, conssistant with geography.
Reference- Britt, Chure, Currie, Holmes, Theurer and Scheetz, 2021. A new deinonychosaurian theropod from the Mid-Cretaceous (Albian) Mussentuchit Member of the Cedar Mountain Formation in Dinosaur National Monument, northeastern Utah, USA. The Society of Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual Meeting. 65-66

undescribed Caenagnathidae (Fiorillo, 1989)
Late Campanian, Late Cretaceous
Judith River Formation, Montana
Material- two specimens
Comments- Referred to Chirostenotes, but may be Caenagnathus or Citipes instead.
Reference- Fiorillo, 1989. The vertebrate fauna from the Judith River Formation (Late Cretaceous) of Wheatland and Golden Valley Counties, Montana. The Mosasaur. 4, 127-142.

Caenagnathidae indet. (Longrich, Barnes, Clark and Millar, 2013)
Late Campanian, Late Cretaceous
Aguja Formation, Texas, US
Material- (TMM 42335-40) caudal centrum
(TMM 42920-2) incomplete manual ungual II (~80 mm)
(TMM 43057-354) distal metatarsal III (~15.5 mm trans)
(TMM 43057-357) pedal ungual
(UTEP B38 L-3) femur (350 mm)
Comments- Longrich et al. (2013) refer TMM 42920-2 and UTEP B38 L-3 to Chirostenotes sp. based on medium size, but they might also be juvenile Caenagnathus so are assigned to Caenagnathidae indet. here. They assign TMM 42335-40, TMM 43057-354 and TMM 43057-357 to their new taxon Leptorhynchos gaddisi based on small size, but the latter is here considered indeterminate and possibly juvenile. These isolated small postcrania could be juvenile Chirostenotes or adult Citipes (provisionally known from the formation based on tarsometatarsal fragment TMM 43057-36).
Reference- Longrich, Barnes, Clark and Millar, 2013. Caenagnathidae from the Upper Campanian Aguja Formation of west Texas, and a revision of the Caenagnathinae. Bulletin of the Peabody Museum of Natural History. 54(1), 23-49.

Caenagnathidae indet. (Yun and Funston, 2021)
Middle-Late Campanian, Late Cretaceous
Fales Rocks Locality (UW V-81006), Mesaverde Formation, Wyoming, US
Material- (UW 44439) incomplete metatarsal I (~32.7 mm)
Comments- Yun and Funston (2021) described this and concluded "UW 44439 is particularly similar to Anzu wyliei and CMN 8538, and contrasts in several features with Chirostenotes pergracilis", namely "the metatarsal is teardrop-shaped in extensor (dorsal) view, owing to the proximally-tapering shaft and a rounded distal condyle" and "the distal articulation surface of UW 44439 is nearly spherical in dorsal view." They placed it in Caenagnathidae indet..
Reference- Yun and Funston, 2021. A caenagnathid oviraptorosaur metatarsal from the Mesaverde Formation (Campanian) of Wyoming, USA. Vertebrate Anatomy Morphology Palaeontology. 9, 105-115.

undescribed caenagnathid (Hunt-Foster and Foster, 2015)
Late Campanian-Early Maastrichtian, Late Cretaceous
Williams Fork Formation, Colorado, US
Material- humerus
Comments- Hunt-Foster and Foster (2015) state this resembles Microvenator more than Anzu.
Reference- Hunt-Foster and Foster, 2015. First occurrence of an oviraptorosaur (Theropoda: Maniraptora) from the Mesaverde Group (Williams Fork Formation) of northwestern Colorado. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 148.

Caenagnathidae indet. (Jasinski, Sullivan and Lucas, 2011)
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Ojo Alamo Formation, New Mexico, US
Material- (SMP VP-2172) incomplete pedal ungual
Reference- Jasinski, Sullivan and Lucas, 2011. Taxonomic composition of the Alamo Wash local fauna from the Upper Cretaceous Ojo Alamo Formation (Naashoibito Member), San Juan Basin, New Mexico. In Sullivan, Lucas and Spielmann (eds.). Fossil Record 3. New Mexico Museum of Natural History and Science Bulletin. 53, 216-271.

Caenagnathidae indet. (Buckley, 2002)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US
Material- (NS.31996.114H) distal metatarsal II (~216 mm) (Buckley, 2002)
(NS.32001.017B) distal metatarsal II (Buckley, 2002)
(TMP 1996.005.0012) distal metatarsal III (Funston et al., 2016)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US
metatarsal (Anonymous, 1997)
material (DePalma, 2010)
Comments- Anonymous (1997) reported a Chirostenotes metatarsal associated with Tyrannosaurus specimen FMNH PR2081.
Buckley (2002) referred two distal metatarsals II to Citipes (her Elmisaurus elegans) because of their longitudinal ridge which would have made the posterior metatarsus surface deeply concave as in Elmisaurus and Citipes. However, these are the size of large Chirostenotes specimens.
DePalma (2010) referred material to Chirostenotes and Caenagnathus.
Funston et al. (2016) figured a tiny distal metatarsal III (TMP 1996.005.0012) as Leptorhynchos elegans, claiming well-defined cruciate ridges as in Elmisaurus and Citipes, and that the distal articular surface was taller than wide as in Citipes. However, all verified specimens of Citipes have surfaces wider than tall as in Elmisaurus. While this and the other three figured metatarsals (TMP 1995.403.0010, 1984.163.0036 and 1986.036.0186) have surfaces taller than wide, that has not been shown to be taxonomically informative. Note while the figure caption states it and the other figured metatarsals are "from the Upper Campanian Dinosaur Park Formation, Dinosaur Provincial Park, Alberta, Canada" the TMP online catalogue shows this specimen is from the Hell Creek Formation of Montana. While these metatarsi may be Elmisaurus or Citipes, the large coeval Anzu has not had its distal metatarsals described yet (though TMP 1996.005.0012 would have to be a juvenile if referred to it).
References- Anonymous, 1997. Tyrannosaurus rex; A Highly Important and Virtually Complete Fossil Skeleton. Sotheby’s. 56 pp.
Buckley, 2002. New material of Elmisaurus (Theropoda, Elmisauridae) from the Late Cretaceous Hell Creek Formation of southeastern Montana. Journal of Vertebrate Paleontology. 22(3), 39A.
DePalma, 2010. Geology, taphonomy, and paleoecology of a unique Upper Cretaceous bonebed near the Cretaceous-Tertiary boundary in South Dakota. Masters thesis, University of Kansas. 227 pp.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica Polonica. 61(1), 159-173.

undescribed caenagnathid (Breithaupt, 1994)
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming
Comments- This is in a faunal list as Chirostenotes.
Reference- Breithaupt, 1994. Wyoming Dinosaur Diversity. Forty-Fourth Annual Field Conference-1994. Wyoming Geological Association Guidebook. 101-104.

Caenagnathidae indet. (TMP online)
Late Cretaceous?
Alberta?, Canada?
Material- (TMP 1978.040.0001) metacarpal

Caenagnathidae indet. (TMP online)
Middle-Late Campanian, Late Cretaceous
Belly River Group or Bearpaw Formation, Alberta, Canada
Material- (TMP 1979.014.0799) phalanx
(TMP 1980.016.2095) incomplete pubis (Rhodes, Funston and Currie, 2020)

Caenagnathidae indet. (TMP online)
Middle-Late Campanian, Late Cretaceous
Belly River Group, Alberta, Canada
Material- (TMP 1965.025.0036) phalanx (TMP online)
(TMP 1978.019.0002) vertebra (TMP online, as Chirostenotes)
(TMP 1979.008.0137) dorsal vertebra (TMP online, as Chirostenotes)
(TMP 1979.008.0138) sacral vertebra (TMP online, as Chirostenotes)
(TMP 1979.008.0339) manual phalanx (TMP online)
(TMP 1979.008.0366) vertebra (TMP online, as Chirostenotes)
(TMP 1979.008.0743) vertebra (TMP online, as Chirostenotes)
(TMP 1979.008.0823) metatarsal (TMP online, as Chirostenotes)
(TMP 1980.016.0810) cranial element (TMP online)
(UALVP 49064) dorsal vertebra (UALVP online)

Caenagnathidae indet. (Currie, 1992)
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada
Material- (TMP 1966.011.0051) vertebra (TMP online, as Chirostenotes)
(TMP 1967.013.0040) phalanx (TMP online, as Chirostenotes)
(TMP 1967.014.0031) ungual (TMP online, as Chirostenotes)
(TMP 1967.016.0038) ungual (TMP online, as Chirostenotes)
(TMP 1967.017.0035) metatarsal (TMP online, as Chirostenotes)
(TMP 1967.017.0038) ungual (TMP online, as Chirostenotes)
(TMP 1967.019.0147) pedal ungual (TMP online, as Chirostenotes)
(TMP 1967.020.0251) caudal vertebra (TMP online, as Chirostenotes)
(TMP 1973.023.0002) ungual (TMP online, as Chirostenotes)
(TMP 1975.011.0033) cervical vertebra (Sues, 1997)
(TMP 1979.008.0799) caudal vertebra (TMP online, as Chirostenotes)
(TMP 1979.014.0508) pedal phalanx (TMP online, as Chirostenotes)
(TMP 1979.014.0902) pedal phalanx (TMP online, as Chirostenotes)
(TMP 1979.014.0923) caudal vertebra (TMP online, as Chirostenotes)
(TMP 1980.008.0281) pedal ungual (TMP online, as Chirostenotes)
(TMP 1980.016.0556) dorsal vertebra (TMP online, as Chirostenotes)
(TMP 1980.016.0557) sacral vertebra (TMP online, as Chirostenotes)
(TMP 1980.016.0578) pedal phalanx (TMP online, as Chirostenotes)
(TMP 1980.016.0994) metatarsal (TMP online, as Chirostenotes)
(TMP 1980.016.1095) pedal phalanx (TMP online, as Caenagnathus)
(TMP 1980.016.1130) sacral vertebra (TMP online, as Chirostenotes)
(TMP 1980.016.1133) ungual (TMP online, as Chirostenotes)
(TMP 1980.016.1481) metatarsal (TMP online, as Chirostenotes)
(TMP 1980.016.1503) sacrum (Funston, 2020)
(TMP 1980.020.0184) ungual (TMP online, as Chirostenotes)
(TMP 1981.019.0252) parietals (Currie, 1992)
(TMP 1981.019.0285) sacrum (Funston, 2020)
(TMP 1982.016.0275) distal pubis, phalanx (Rhodes, Funston and Currie, 2020)
(TMP 1984.163.0102) sacrum (Funston, 2020)
(TMP 1984.167.0044) sacrum (Makovicky, 1995)
?(TMP 1985.065.0001) partial tibiotarsus (Atkins-Weltman, Simon, Woodward, Funston and Snively, 2024)
(TMP 1989.036.0109) caudal vertebra (Makovicky, 1995)
(TMP 1991.036.0146) caudal vertebra (Makovicky, 1995)
(TMP 1992.036.0053) proximal caudal vertebra (31.4 mm) (Currie et al., 1993)
(TMP 1994.012.0603) pubes (Rhodes, Funston and Currie, 2020)
(TMP 1995.403.0010) distal metatarsal III (Funston, Currie and Burns, 2016)
(TMP 1998.093.0013) posterior ilium (Rhodes, Funston and Currie, 2020)
(TMP 2001.012.0216) distal quadrate (Funston, 2020)
(TMP 2002.012.0103) partial ilium (Rhodes, Funston and Currie, 2020)
Comments- Only specimens through 1980 have been added from the TMP online catalogue so far. These are often referred to Caenagnathus or Chirostenotes in the catalogue, but are retained as Caenagnathidae here as differences between these geneera have been contentious and only recently resolved.
TMP 1981.019.0252 and 1992.036.0053 were referred to Caenagnathus sp. by Currie (1992) and Currie et al. (1993) respectively. Sues (1997) referred TMP 1975.011.0033 (misidentified as a caudal on the TMP online catalog) to Chirostenotes, while Makovicky (1995) referred TMP 1984.167.0044, 1989.036.0109 and 1991.036.0146 to that genus. The latter material did not have locality information listed, but is probably from the Dinosaur Park Formation of Alberta. TMP 1995.403.0010 is figured by Funston et al. (2016) as Leptorhynchus elegans (now Citipes), but seems too large. It may be Chirostenotes or young Caenagnathus instead. Funston (2019) describes and figures partial ilium TMP 1998.093.0013, pubes TMP 1994.012.0603 and pubis TMP 1980.016.2095. The pubes differ from the larger one referred to Caenagnathus, so are believed by Funston to be either Chirostenotes or Citipes. Atkins-Weltman et al. (2024) mention "an undescribed partial tibiotarsus" with astragalocalcaneum fused to the tibia as in Eoneophron "possibly also from a caenagnathid, from the Dinosaur Park Formation of Alberta (TMP 1985.065.0001; GFF pers. obs.). The latter specimen (TMP 1985.065.0001) is catalogued as an ornithomimid, but shares several features with caenagnathids, including a semi-circular cross-section of the shaft and the absence of a groove or ridge for the fibula (which sits lateral to the tibia throughout its length in oviraptorosaurs, as opposed to other theropods including ornithomimids)."
Reference- Currie, 1992. Saurischian dinosaurs of the Late Cretaceous of Asia and North America. In Mateer and Chen (eds.). Aspects of Nonmarine Cretaceous Geology. China Ocean Press. 237-249.
Currie, Godfrey and Nessov, 1993 (published 1994). New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30(10), 2255-2272.
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria (Dinosauria: Theropoda). Masters thesis, Copenhagen University. [pp]
Sues, 1997. On Chirostenotes, a Late Cretaceous oviraptorosaur (Dinosauria: Theropoda) from western North America. Journal of Vertebrate Paleontology. 17(4), 698-716.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica Polonica. 61(1), 159-173.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria (Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Funston, 2020. Caenagnathids of the Dinosaur Park Formation (Campanian) of Alberta, Canada: Anatomy, osteohistology, taxonomy, and evolution. Vertebrate Anatomy Morphology Palaeontology. 8, 105-153.
Rhodes, Funston and Currie, 2020. New material reveals the pelvic morphology of Caenagnathidae (Theropoda, Oviraptorosauria). Cretaceous Research. 114, 104521.
Atkins-Weltman, Simon, Woodward, Funston and Snively, 2024. A new oviraptorosaur (Dinosauria: Theropoda) from the end-Maastrichtian Hell Creek Formation of North America. PLoS ONE. 19(1): e0294901.

Caenagnathidae indet. (Eberth, Evans, Brinkman, Therrien, Tanke and Russell, 2013)
Middle Maastrichtian, Late Cretaceous
Tolman Member of Horseshoe Canyon Formation, Alberta, Canada
(UALVP 57349) (1 year old juvenile, ~8.0 kg) incomplete tibia (~210 mm) (Funston and Currie, 2018)
Early Maastrichtian, Late Cretaceous
boundary of Morrin and Tolman Members of Horseshoe Canyon Formation, Alberta, Canada
(WL-156) multiple elements (Eberth et al., 2013)
Comments- Eberth et al. (2013) lists WL-156 as a possible Epichirostenotes specimen, though further details including its whereabouts given the subsequent death of Wann Langston ('WL' stands for his personal collection) are uncertain.
UALVP 57349 is a juvenile tibia found on July 23 2016 and described by Funston and Currie (2018) as Caenagnathidae indet.. While the two recognized Horseshow Canyon caenagnathid taxa preserve tibiae, only the proximal end is exposed in Apatoraptor while only the distal end is preserved in Epichirostenotes. UALVP 57349 is indistinguishable from either, given its mostly broken proximal end and Epichirostenotes' unillustrated and briefly described distal end. The widely expanded distal end (236% of shaft width) is most similar to Elmisaurus (184%) as opposed to Anzu (158%), Chirostenotes (161%) or Citipes (132%).
References- Eberth, Evans, Brinkman, Therrien, Tanke and Russell, 2013. Dinosaur biostratigraphy of the Edmonton Group (Upper Cretaceous), Alberta, Canada: Evidence for climate influence. Canadian Journal of Earth Sciences. 50(7), 701-726.
Funston and Currie, 2018. A small caenagnathid tibia from the Horseshoe Canyon Formation (Maastrichtian): Implications for growth and lifestyle in oviraptorosaurs. Cretaceous Research. 92, 220-230.

undescribed caenagnathid (Ryan and Russell, 2001)
Late Maastrichtian, Late Cretaceous
Scollard Formation, Alberta, Canada
Material- (RTMP 86.207.17) partial cervical vertebra
Comments- This was cited as Segnosauridae indet. by Ryan and Russell (2001) in a faunal list, but Cullen et al. (2020) found it "to be most similar in morphology to caenagnathids."
Reference- Ryan and Russell, 2001. Dinosaurs of Alberta (exclusive of Aves). In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life. Indiana University Press. 279-297.
Cullen, Larson, Zanno, Currie and Evans, 2020. Theropod biodiversity patterns in the Dinosaur Park Formation (Late Cretaceous: Campanian) of Alberta revealed through morphometrics and biostratigraphy. The Society of Vertebrate Paleontology 80th Annual Meeting, Conference Program. 115.

unnamed Caenagnathidae (Gilmore, 1924)
Maastrichtian, Late Cretaceous
Frenchman Formation, Saskatchewan, Canada
Material- (CMN 346) manual ungual I (~130 mm on curve) (Bell, Currie and Russell, 2015)
(CMN 8504) dorsal centrum (44 mm), three caudal centra (22-25 mm) (Gilmore, 1924)
(RSM P2161.1) metatarsal II (~211 mm) (Funston, Currie and Burns, 2016)
(TMM 41395-1) manual ungual II (Bell, Currie and Russell, 2015)
Comments- The centra were described by Gilmore (1924) as distinct from other coelurosaurs known at the time, though possibly referrable to Chirostenotes or Dromaeosauridae (neither of which were then known from vertebrae). Currie et al. (1993) noted the caudals belonged to an oviraptorosaur, referring them to Caenagnathus sp.. Bell et al. (2015) described two manual unguals as cf. Anzu, noting they are comparable in size and age but that CMN 346 has a deeper concavity between the flexor tubercle and articular surface than that genus. Elmisaurus sp. is also present in this formation based on RSM P2600.1. Funston et al. (2016) describe metatarsal II RSM P2161.1 as Leptorhynchos sp., but its large size and unfused distal tarsal are more like Chirostenotes or Caenagnathus. It lacks the large posterior protuberance on the proximal end seen in Caenagnathus, and has a more prominent posteromedial ridge as in Elmisaurus and Citipes.
References- Gilmore, 1924. A new coelurid dinosaur from the Belly River Cretaceous Alberta. Canada Geological Survey, Bulletin n. 38, geological series 43, 1-13.
Tokaryak, 1990. It was here a minute ago. The Saskatchewan Archaeological Society Newsletter. 11(2), 44-45.
Currie, Godfrey and Nessov, 1993 (published 1994). New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30(10), 2255-2272.
Bell, Currie and Russell, 2015 (online 2014). Large caenagnathids (Dinosauria, Oviraptorosauria) from the uppermost Cretaceous of western Canada. Cretaceous Research. 52, 101-107.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica Polonica. 61(1), 159-173.

unnamed caenagnathid (Nessov and Khisarova, 1988)
Santonian, Late Cretaceous
Bostobe Formation, Kazakhstan
Material- dentary
Comments- This was first described as a turtle (Nessov and Kisarova, 1988), but later identified as a caenagnathid and relative of Caenagnathasia by Currie et al. (1993).
References- Nessov and Khisarova, 1988. New data on vertebrates from the Late Cretaceous of Shakh-Shakh and Baybolat (northeastern Aral region). In Material on the history of the fauna and flora of Kazakhstan, Vol. 10. Academy of Sciences of Kazakhstan, Alma Ata. 5-14.
Currie, Godfrey and Nessov, 1993 (published 1994). New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30(10), 2255-2272.

undescribed caenagnathid (Manabe and Barrett, 2000)
Valanginian-Hauterivian, Early Cretaceous
Kuwajima Formation of the Tetori Group, Japan
Material- (SBEI-167) manual ungual
Comments- Referred to the oviraptorosaur-therizinosaur clade by Manabe et al. (2000), it provisionally resembles caenagnathids most closely.
References- Barrett and Manabe, 2000. The dinosaur fauna from the Earliest Cretaceous Tetori Group of Central Honshu, Japan. Journal of Vertebrate Paleontology. 20(3), 28A-29A.
Manabe and Barrett, 2000. Dinosaurs. In Matsuoka (ed.). Fossils of the Kuwajima "Kaseki-kabe" (fossil-bluff). Scientific report on a Neocomian (Early Cretaceous) fossil assemblage from the Kuwajima Formation, Tetori Group, Shiramine, Ishikawa, Japan. Shiramine Village Board of Education, Japan. 93-98.
Manabe, Barrett and Isaji, 2000. A refugium for relicts? Nature. 404, 953-954.
Matsuoka, Kusuhashi, Takada and Setoguchi, 2002. A clue to the Neocomian vertebrate fauna: Initial results from the Kuwajima 'Kaseki-kabe' (Tetori Group) in Shiramine, Ishikawa, central Japan. Memoirs of the Faculty of Science, Kyoto University, Series of Geology and Mineralogy. 59(1), 33-45.

Microvenator Ostrom, 1970
= "Megadontosaurus" Ostrom, 1970
M. celer Ostrom, 1970
= "Megadontosaurus ferox" Brown vide Chure and McIntosh, 1989
Mid-Late Albian, Early Cretaceous
Himes Member, Cloverly Formation, Montana, US
Holotype- (AMNH 3041 in part) (~3.7 kg; juvenile) lacrimals(?) (one lost), quadrates(?) (lost), dentary, prearticular (?) (lost), axis (13.9 mm), anterior cervical neural arch, cervical centrum (15.1 mm), cervical centrum (15.4 mm), cervical centrum (15+ mm), two cervical neural arches, two partial cervical ribs, three anterior dorsal neural arches, anterior dorsal centrum (13.5 mm), anterior dorsal centrum (13.8 mm), nine posterior dorsal neural arches, posterior dorsal centrum (14.2+ mm), posterior dorsal centrum (13.8 mm), posterior dorsal centrum (13.9 mm), posterior dorsal centrum (14.1 mm), posterior dorsal centrum (13.7 mm), posterior dorsal centrum (13.1 mm), two partial dorsal ribs, mid sacral centrum (15.1 mm), caudal centrum (11.5 mm), caudal centrum (9.6+ mm), caudal centrum (8.7 mm), caudal centrum (6.1+ mm), caudal centrum (9.9+ mm), caudal centrum (9.8 mm), caudal centrum (9.7 mm), caudal centrum (9.5 mm), caudal centrum (10.1 mm), three caudal neural arches, coracoid, humerus (81.5 mm), radius, ulna (78 mm), ulnar fragment, metacarpal I (12.5 mm), phalanx I-1 (34 mm), manual ungual I (25 mm), manual ungual III (11 mm), distal phalanx, partial ilia (~110 mm), pubes (108.6 mm), fragmentary ischia, femora (124 mm), tibia (157 mm), proximal fibula, astragalus (22.5 mm wide, 42 mm high), metatarsal I (11.8+ mm), pedal ungual I (11.9 mm)
Diagnosis- (after Makovicky and Sues, 1998) dorsal and caudal centra distinctly wider than high; accessory trochanter at base of the anterior trochanter of the femur.
Other diagnoses- (after Ostrom, 1970) very small (approximately one half to two thirds the size of Ornitholestes or Coelurus), delicately built; hollow, thin-walled vertebrae and limb bones. cervicals without neural spines and with double pleurocels; dorsal neural arches low and highly sculpted, neural spines low and rectangular, postzygapophyses far behind posterior border of centrum; pubis profile concave anteriorly, distal extremities only moderately expanded; femur with short but prominent anterior trochanter and a depression at site of fourth trochanter; astragalus with very high and broad ascending process.
Comments- The holotype was collected in 1933 but not described until 1970 (Ostrom, 1970). As noted by Ostrom, "associated with the type specimen were 25 teeth of the Deinonychus type which Barnum Brown believed belonged to this specimen." "Because of the extremely large size of the associated teeth, this specimen has been known informally in conversation by the name "Megadontosaurus"" As Chure and McIntosh (1989) stated, Brown proposed "Megadontosaurus ferox" with the intended holotype AMNH 3041, but "never got around to describing these animals, and it was left to John Ostrom to describe and name them many years later in 1970, based in many cases on much better and more diagnostic material which he had collected." "None of Brown's names were ever published (until Glut 1972 [for "Peltosaurus"]) and it does no harm to mention them here, as Ostrom's names are well established taxa having full priority. Brown's names, however, were used in public lectures, and for a time on some exhibited specimens. In addition, they were written on photographs and preliminary skeletal reconstructions made under Brown's direction." Ostrom writes "without excveption, the 25 associated teeth duplicate in size and form the teeth found with the type of Deinonychus antirrhopus", including the high DSDI. He referred them to that taxon, and noted "they are still catalogued with the Microvenator remains under AMNH 3041."
The tooth (YPM 5366) questionably referred to Microvenator by Ostrom is unlikely to belong to an oviraptorosaur, as it is plesiomorphically recurved and ziphodont, so is referred to Tetanurae indet. here. The same is true of another tooth (MOR coll.) referred to Microvenator by Maxwell (1993). Yun and Funston (2021) figured the first metatarsal in detail.
This genus has a controversial placement in recent analyses, from being a caudipterid in Senter et al. (2012), avimimid in Lee et al. (2014), sister to caenagnathoids in Brusatte et al. (2014) and Foth et al. (2014), and basal caenagnathid in Funston and Currie (2016). Hartman et al. (2019) recovered it as a basal oviraptorid, but with the addition of more taxa it moved to the first branching caenagnathid.
References- Ostrom, 1970. Stratigraphy and paleontology of the Cloverly Formation (Lower Cretaceous) of the Bighorn Basin area, Wyoming and Montana. Peabody Museum Bulletin. 35, 234 pp.
Chure and McIntosh, 1989. A Bibliography of the Dinosauria (Exclusive of the Aves) 1677-1986. Museum of Western Colorado Paleontology Series #1. 226 pp.
Maxwell, 1993. Neonate dinosaur remains and dinosaur eggshell from the Lower Cretaceous Cloverly Formation of Montana. Journal of Vertebrate Paleontology. 13(3), 48A.
Makovicky and Sues, 1997. A reappraisal of the phylogenetic affinities of Microvenator celer (Theropoda: Dinosauria) from the Cloverly Formation. Journal of Vertebrate Paleontology. 17(3), 62A.
Makovicky and Sues, 1998. Anatomy and phylogenetic relationships of the theropod dinosaur Microvenator celer from the Lower Cretaceous of Montana. American Museum Novitates. 3240, 27 pp.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid tail. PLoS ONE. 7(5), e36790.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan culminated in rapid rates of evolution across the dinosaur-bird transition. Current Biology. 24(20), 2386-2392.
Foth, Tischlinger and Rauhut, 2014. New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers. Nature. 511, 79-82.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247.
Yun and Funston, 2021. A caenagnathid oviraptorosaur metatarsal from the Mesaverde Formation (Campanian) of Wyoming, USA. Vertebrate Anatomy Morphology Palaeontology. 9, 105-115.

Caenagnathidae sensu Sues, 1997
Definition- (Chirostenotes pergracilis + Chirostenotes elegans + Elmisaurus rarus + Caenagnathasia martinsoni + BHM 2033)
Comments- Phylogeny and taxonomy within this group have been incredibly contentious, along with which elements can be referred to each species. The listed diagnoses are an exception to the norm for this website in that they only compare to this group of caenagnathids and not other taxa on this page like Microvenator or oviraptorids, even if avimimids are correctly nested within the clade.

Leptorhynchos Longrich, Barnes, Clark and Millar, 2013b
= "Leptorhynchos" Longrich, Barnes, Clark and Millar, 2013a
L. gaddisi Longrich, Barnes, Clark and Millar, 2013b
= "Leptorhynchos gaddisi" Longrich, Barnes, Clark and Millar, 2013a
Late Campanian, Late Cretaceous
Aguja Formation, Texas, US
Holotype- (TMM 45920-1) (juvenile?) dentaries (symph 24.45 mm)
Diagnosis- Provisionally indeterminate relative to Chirostenotes and Elmisaurus.
Other diagnoses- Longrich et al. (2013a) diagnosed Leptorhynchos gaddisi based on comparison with TMP 1992.036.0390, a specimen they referred to elegans (as Leptorhynchos elegans, now separated as Citipes elegans). However, that specimen is here considered Chirostenotes pergracilis based on its large size and juvenile features. In turn, they diagnosed the genus Leptorhynchos based on comparison with TMP 1990.056.0006, another mandible of Chirostenotes pergracilis. Of characters listed as diagnostic for Leptorhynchos, the shortness or depth of the dentary and dentary contribution to the dorsal margin of the external mandibular fenestra cannot be evaluated in the type as the posterior section is missing. An upturned dentary tip and four symphyseal ridges are also present in Chirostenotes (e.g. complete mandibles TMP 2001.012.0012). "Dentaries strongly divergent, with posterior rami of dentaries forming an angle of roughly 35 degrees in dorsal view" is not true in the L. gaddisi holotype. Indeed, the diagnosis of L. gaddisi lists "lateral occlusal margins of the dentaries not as strongly divergent in dorsal view, with the tip of the beak being narrower and more spoon-shaped in dorsal view", which is true as they show an angle of about 20 degrees. Both this and the "short caudal symphyseal shelf" fall within the range of variation of Chirostenotes (again compare to TMP 2001.012.0012 which has a ~20 degree angle and a shorter symphysis than gaddisi's holotype). Of the remaining two listed characters for L. gaddisi, "a strongly rounded anteroventral margin of the symphysis" is indistinguishable from some Chirostenotes (TMP 1992.036.0390) and "a more anteriorly projecting tip of the beak" than 1992.036.0390 is true in other Chirostenotes jaws like 2001.012.0012. Thus all suggested characters fall within the range of variation of Chirostenotes pergracilis. The size is relatively small but within the range of known juvenile Chirostenotes, plus Funston et al. (2020) showed dentary fusion occurs even in those juveniles, so this does not suggest "that the animal was mature" as Longrich et al. claim. Intriguingly, the Leptorhynchos holotype seems to show fibrous external bone texture (Longrich et al.'s Fig. 8D below the label) and a poorly developed m. genioglossus attachment, both of which Funston et al. suggested could be associated with immaturity. Thus it could easily be a juvenile of a Chirostenotes-sized taxon. As it cannot presently be distinguished from Chirostenotes or the cf. Elmisaurus rarus dentaries IGM 102/107, it is here considered an indeterminate caenagnathid.
Comments- Some of this material was first reported as Caenagnathidae indet. by Longrich et al. (2010). While Longrich et al. (2013a) intended to name Leptorhynchos gaddisi, they did not specify a type species, making the correction Longrich et al. (2013b) necessary to specify gaddisi as the type species and make the names official.
Longrich et al. (2013a) proposed this species is sister to elegans, both being closer to Chirostenotes than to Elmisaurus rarus. Their phylogenetic analysis would seem to support this, but Elmisaurus rarus' basal position is due to their figure 14 being an Adams consensus tree. Because rarus lacks described mandibular material, it cannot be placed precisely compared to Caenagnathasia and their Caenagnathus OTUs, so will have a position at the base of the largest clade it can belong to in an Adams consensus tree. When their analysis is rerun to a posteriori exclude taxa incomparable to Elmisaurus rarus, it is in a trichotomy with Chirostenotes and elegans. Further, their analysis miscodes rarus as lacking metatarsal fusion and excludes the other five characters suggested by Currie (1989) to unite rarus and elegans to the exclusion of pergracilis.
Longrich et al. do claim elegans is more similar to Chirostenotes in one way. They state "Longrich (2008a) tentatively placed [elegans] in Chirostenotes, because the third metatarsal has an anteroposteriorly flattened shaft that is concave and broadly exposed on the posterior of the metatarsus (Currie 1989). This is a derived feature found in Chirostenotes (Currie and Russell 1988) but not Elmisaurus (Osmólska 1981)." This seems related to their new character 205- "Metatarsal III with an ovoid or subtriangular cross section (0) or anteroposteriorly flattened, with a concave posterior surface (1). Primitively in theropods the third metatarsal has an ovoid cross section, or a triangular cross section in arctometatarsalian forms. This condition is retained in most oviraptorosaurs, including the basal caenagnathid Elmisaurus rarus. In Caenagnathinae, the third metatarsal is anteroposteriorly compressed." Yet the posterior transverse exposure of proximal metatarsal III proximally seems intermediate in E. rarus' holotype compared to the two specimens of elegans. More distally, Currie's (1989) figure 2P section indicates the posterior exposure is narrow as in E. rarus. Currie's (1989) figure 2Q shows E. rarus has a concave posterior metatarsal III surface as well. As for shape, the main issue seems to be the use of different proximodistal points along the bone. Currie and Russell (1988) state in Chirostenotes pergracilis "The proximal end, viewed dorsally, is diamond shaped, tapering both anteriorly (between the contact of metatarsals II and IV) and posteriorly. Its major horizontal axis, 17.5 mm long, is anteroposterior in orientation and thins backwards." Sternberg (1932) also states the Macrophalangia holotype (possibly Caenagnathus collinsi according to Longrich et al.) has a transversely compressed proximal end. The proximal ends of E. rarus' and elegans' metatarsal III are fused too well with surrounding bones to compare. Once C. pergracilis' anterior surface is exposed, "the bone twists until the medial surface is facing anteriorly" and "the anterior edge has broadened out to 7.5 mm to separate the adjacent metatarsals and is triangular in section" (Currie and Russell, 1988). elegans' holotype is broken at about this same point and also shows a triangular section. Note a triangular section is what Longrich et al. are claiming caenagnathines don't have, though it exists in both pergracilis and elegans. Currie (1989) was also wrong in comparing the proximal diamond shape of Chirostenotes pergracilis with the more distal triangular shape of elegans, and none of these areas have been described in E. rarus. At two-thirds down in Chirostenotes, "In section, a shallow concave surface faces posteriorly at this level, while slightly concave surfaces face posteromedially and posterolaterally for contact with the adjacent metatarsals." This appears similar to E. rarus from what the anterior and posterior views suggest, and matches how Snively (2000) described the specimen. It also matches the cross section of elegans illustrated by Currie (1989- fig. 2P). While Currie's figure 2Q of Elmisaurus rarus would suggest a slightly different shape where the articular surfaces are smaller and that for metatarsal IV doesn't angle posteromedially, the narrow anterior exposure of metatarsal III means it must have been taken more proximally, probably about halfway down considering the ratio between anterior and posterior exposure of metatarsal III. This leaves anteroposterior compression, which varies throughout the bone in elegans at least. Distally it's transversely compressed but proximally it's anteroposteriorly compressed. The E. rarus section which is probably intermediate in position is also intermediate in compression, being slightly transversely compressed. If Chirostenotes specimens are any indication, the bone switches back to transversely compressed at its proximal tip. So there are actually no valid described differences in metatarsal III sectional shape between Chirostenotes and any Elmisaurus species in the primary literature.
As discussed above under Other diagnoses, the Leptorhynchos gaddisi holotype is indeterminate and possibly juvenile. None of the referred material is diagnostic, but the small and fused distal tarsal and metatarsal IV (TMM 43057-36) are like Citipes and Elmisaurus. If anything, this suggests the large Aguja ungual and femur (TMM 42920-2 and UTEP B38 L-3) described by Longrich et al. might belong to adult Leptorhynchos/Chirostenotes, while TMM 43057-36 is a separate smaller species, perhaps Citipes. The small caudal centrum (TMM 42335-40), distal metatarsal III (TMM 43057-354) and pedal ungual (TMM 43057-357) are here placed in Caenagnathidae indet., as they could be juveniles, while the proximal partial tarsometatarsus is provisionally referred to the contemporeneous Citipes elegans.
References- Sternberg, 1932. Two new theropod dinosaurs from the Belly River Formation of Alberta. Canadian Field-Naturalist. 46(5), 99-105.
Currie and Russell, 1988. Osteology and relationships of Chirostenotes pergracilis (Saurischia, Theropoda) from the Judith River (Oldman) Formation of Alberta, Canada. Canadian Journal of Earth Sciences. 25(7), 972-986.
Currie, 1989. The first records of Elmisaurus (Saurischia, Theropoda) from North America. Canadian Journal of Earth Sciences. 26(6), 1319-1324.
Snively, 2000. Functional morphology of the tyrannosaund arctometatarsus. Masters Thesis, University of Calgary. 273 pp.
Longrich, Sankey and Tanke, 2010. Texacephale langstoni, a new genus of pachycephalosaurid (Dinosauria: Ornithischia) from the upper Campanian Aguja Formation, southern Texas, USA. Cretaceous Research. 31, 274-284.
Longrich, Barnes, Clark and Millar, 2013a. Caenagnathidae from the Upper Campanian Aguja Formation of west Texas, and a revision of the Caenagnathinae. Bulletin of the Peabody Museum of Natural History. 54(1), 23-49.
Longrich, Barnes, Clark and Millar, 2013b. Correction to "Caenagnathidae from the Upper Campanian Aguja Formation of west Texas, and a revision of the Caenagnathinae". Bulletin of the Peabody Museum of Natural History. 54(2), 263-264.
Funston, Wilkinson, Simon, Leblanc, Wosik and Currie, 2020 (online 2019). Histology of caenagnathid (Theropoda, Oviraptorosauria) dentaries and implications for development, ontogenetic edentulism, and taxonomy. The Anatomical Record. 303(4), 918-934.

Ojoraptorsaurus Sullivan, Jasinski and van Tomme, 2011
O. boerei Sullivan, Jasinski and van Tomme, 2011
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Ojo Alamo Formation, New Mexico, US
Holotype- (SMP VP-1458) (~1.8-2.1 m) incomplete pubes (~344 mm)
Diagnosis- (after Sullivan et al., 2011) iliac peduncle articular surface of pubes narrower anteriorly than posteriorly (also in Microvenator, Caenagnathus, Chirostenotes and Chirostenotes/Citipes TMP 1994.012.0603; unknown in other caenagnathids except Epichirostenotes); proximomedial pubic fossa recessed distally from acetabular rim at least one third the narrowest anteroposterior width of the ilial peduncle (unknown in other caenagnathids except Microvenator, Anzu, Epichirostenotes, Chirostenotes/Citipes TMP 1980.016.2095 and Elmisaurus?); oval depression on the anterodorsal surface of pubic boot (also in Microvenator and Epichirostenotes; unknown in other caenagnathids except Anzu and Chirostenotes/Citipes TMP 1994.012.0603); pubic shaft strongly convex anteriorly just proximal to pubic boot (unknown in other caenagnathids except Microvenator, Anzu, Caenagnathus, Epichirostenotes?, Chirostenotes/Citipes TMP 1980.016.2095 and TMP 1994.012.0603 and Elmisaurus?).
Other diagnoses- Sullivan et al. (2011) also listed differences from Epichirostenotes "in being 20% smaller and relatively more robust, lacking a dorsal suture on the pubic boot, and having a significantly lower pubic apron/pubis length ratio." However, all other caenagnathid pubes are more robust than Epichirostenotes' seems to be, perhaps due to the amount of crushing in that taxon's holotype. The small size difference with undescribed histology in each specimen means little, and that "there is a trace of a suture" on the dorsal surface of Epichirostenotes' pubis is not a strong indicator of a less mature specimen. The apron/pubis ratio (supposedly 38% vs. 49%) actually refers to the distance from the distal interpubic foramen tip to the proximomedial corner of the apron, based on its stated length in Ojoraptorsaurus. Yet this is 39% in Epichirostenotes according to Sues' figure, comparable to Anzu's 38% and Ojoraptorsaurus' estimated 38% (Caenagnathus and Chirostenotes or Citipes TMP 1980.016.2095 are now known to be shorter at 33% and 30% respectively). So it seems Sullivan et al. overestimated the length in Epichirostenotes.
Comments- The holotype was discovered in summer 2002.
The validity of this taxon was initially uncertain, as it was only compared to Nomingia (here an oviraptorid), Epichirostenotes and Anzu by Sullivan et al. (2011) and the distribution of its diagnostic features in these taxa and other oviraptorosaurs was poorly understood. Since then, pubes of Caenagnathus, Chirostenotes or Citipes and Elmisaurus have been described. These show Ojoraptorsaurus to be diagnosable, although the sub-trapezoidal ilial articulation would seem to be a symplesiomorphic difference from Epichirostenotes. The amount of separation of the proximomedial fossa from the acetabulum should be determinable in Caenagnathus and Elmisaurus as well, but is currently undescribed and unfigured.
Reference- Sullivan, Jasinski and van Tomme, 2011. A new caenagnathid Ojoraptorsaurus boerei, n. gen., n. sp. (Dinosauria, Oviraptorosauria), from the Upper Cretaceous Ojo Alamo Formation (Naashoibito Member), San Juan Basin, New Mexico. New Mexico Museum of Natural History and Science Bulletin. 53, 418-428.

Kuszholia Nessov, 1992
?= Caenagnathasia Currie, Godfrey and Nessov, 1993
K. mengi Nessov, 1992
?= Caenagnathasia martinsoni Currie, Godfrey and Nessov, 1993
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Holotype- (ZIN PO 4602) posterior synsacrum (11, 10 mm)
Paratypes- ?(ZIN PO 4623) anterior synsacral centra (13, 10 mm)
?(ZIN PO coll.) sacral vertebrae (~20-30 mm)
Referred- ?(CCMGE 401/12457; holotype of Caenagnathasia martinsoni) anterior dentaries (symph 11.2 mm) (Currie, Godfrey and Nessov, 1993)
?(CCMGE 402/12457; paratype of Caenagnathasia martinsoni) incomplete dentary (symph 8.94 mm) (Currie, Godfrey and Nessov, 1993)
?(CCMGE 479/12457) proximal femur (Nessov, 1995; described by Sues and Averianov, 2015)
?(ZIN PH 802/16) four incomplete fused sacral vertebrae (17, 14, 16.3, 16.4 mm) (Sues and Averianov, 2015)
?(ZIN PH 932/16) incomplete posterior cervical vertebra (17 mm) (Sues and Averianov, 2015)
?(ZIN PH 933/16) incomplete anterior cervical vertebra (19.5 mm) (Sues and Averianov, 2015)
?(ZIN PH 934/16) incomplete dorsal vertebra (16.4 mm) (Sues and Averianov, 2015)
?(ZIN PH 935/16) dorsal vertebra (Sues and Averianov, 2015)
?(ZIN PH 936/16) dorsal vertebra (Sues and Averianov, 2015)
?(ZIN PH 937/16) posterior dorsal centrum (Sues and Averianov, 2015)
?(ZIN PH 2354/16) anterior dentaries (symph 7.5 mm) (Sues and Averianov, 2015)
?(ZIN PO 4603) incomplete anterior cervical vertebra (16 mm) (Nessov, 1992; described by Sues and Averianov, 2015)
?(ZIN PO 5234) partial cervical vertebra (Sues and Averianov, 2015)
? (lost) mandible with teeth(?) (Funston, Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020)
Diagnosis- (after Currie et al., 1993) first anterior occlusal groove much larger than second grooves (when present) (unknown in other caenagnathids except Caenagnathus, Chirostenotes and Elmisaurus); lateral occusal ridges do not meet anteriorly (also in Anzu, Chirostenotes, Elmisaurus and Citipes; unknown in other caenagnathids except Caenagnathus); lingual ridge with taller and sharper edge (the cause of their "lateral groove is narrower and deeper") (also in Caenagnathus; unknown in other caenagnathids except Anzu, Chirostenotes, Elmisaurus and Citipes); defined medial edges to fossae between lingual grooves (unknown in other caenagnathids except Anzu, Caenagnathus, Chirostenotes, Elmisaurus and Citipes); vascular grooves not conspicuous on symphysial shelf (also in Caenagnathus and Elmisaurus; unknown in other caenagnathids except Anzu, Chirostenotes and Citipes); no foramina on the floor of median depression posterior to first anterior occlusal groove (unknown in other caenagnathids except Anzu?, Caenagnathus, Chirostenotes, Elmisaurus and Citipes).
(suggested) large sacral pleurocoels (unknown in other caenagnathids except Chirostenotes, Epichirostenotes and Citipes).
Other diagnoses- None of the characters listed in Nessov's (1992) diagnosis of Kuszholia are very characteristic. Dorsoventrally compressed sacral centra are common in maniraptorans. The robust second to last sacral transverse process is seen in Chirostenotes and Epichirostenotes as well. Deep posterior sacral pleurocoels are common in caenagnathoids, and far from being small, Kuszholia's are large compared to other taxa. The posterior articular surface is not large, being smaller than mid sacral vertebrae, and its concavity is plesiomorphic for theropods. The large postzygapophyses are only notable compared to birds- they are normal for an oviraptorosaur. The sacrals of Chirostenotes also have a ventral median groove which is most pronounced at the junction of centra.
Currie et al. (1993) stated there was no second anterior occlusal groove in Caenagnathasia, but Sues and Averianov (2015) reported it as present in new specimen ZIN PH 2354/16 and showed it is present although small on the right side of the holotype as well. Currie et al. also stated the "fluting on the lingual margin of the occlusal edge is not as distinct in the Asian jaws, and there are no toothlike apical projections on the ridges" unlike Caenagnathus (including Chirostenotes), but this is untrue for ZIN PH 2354/16. They said "there are no tubercles on the midline or at the base of the first lateral occlusal ridge", but Funston et al. (2020a) found that in Chirostenotes "some dentaries lack these nodules altogether (TMP 1990.056.0006; TMP 2001.012. 0012). Therefore, the presence or absence of these rugose patches is probably best explained by individual variation." Indeedd, a tubercle is on the midline in ZIN PH 2354/16.
Comments- Note that while volume 30(10) of the Canadian Journal of Earth Sciences lists its date as October 1993, it was not published until February or March of 1994.
The holotype (ZIN PO 4602) consists of the last two sacral vertebrae and a fragment of the third to last, fused together. The centra are dorsoventrally compressed (anterior articular surface ~66% as tall as wide) with an oval and slightly concave posterior articular surface. The sacrum seems to be slightly concave ventrally and the centrum junctions are expanded both ventrally and laterally. There are small but deep pleurocoels present in each centrum. A median ventral groove is present, which is especially well marked at the centrum junctions. The postzygapophyses of the last sacral protrude markedly past the centrum. The second to last transverse process is long, robust and perpendicular to the sacral long axis, while the last transverse process is about half the length but otherwise similar.
Nessov referred another specimen (ZIN PO 4623) consisting of the first two fused centra of another sacrum. These are similar in being dorsoventrally compressed (posterior articular surface ~63% as tall as wide). The anterior articular surface is kidney-shaped and slightly heterocoelous. It is also similar in being slightly concave, with expanded centrum junctions and deep, oval pleurocoels in each centrum. Ventrally, there is a slight midline groove. This was later referred to Caenagnathasia by Sues and Averianov (2015). Nessov referred to additional isolated vertebrae virtually identical to these, but 2-2.5 times larger. These may belong to larger oviraptorosaurs, therizinosaurs or dromaeosaurids, though they are impossible to evaluate without more information. Finally, he stated strongly pneumatic vertebrae with closed neurocentral sutures (unlike juvenile therizinosaurs or sauropods) could belong to Kuszholia. One is illustrated (ZIN PO 4603), which is an anterior cervical with strongly overhanging prezygapophyses, a large neural canal, an elongate centrum (2.75 times posterior height) which reaches posteriorly past the neural arch, and perhaps a large teardrop shaped pleurocoel. This was later referred to Caenagnathasia by Sues and Averianov (2015).
Nessov and Panteleev (1993) figured and described a partial sacrum they referred to Kuszholia sp. (ZIN PO 4826). Zelenkov and Averianov (2011) stated this differs from Kuszholia in "the absence of a pleurocoel in the posterior vertebra and in the shallow slitlike pleurocoel in the penultimate vertebra", and while the pleurocoel shape appears similar, the absence of a pleurocoel in the last sacral is indeed different. They believe the specimen to be similar to Zanabazar, but I don't see any particular resemblence and refer it to Maniraptora incertae sedis here pending further study. Nessov (1995) later figured a centrum with a slit-like pleurocoel (ZIN PO 467) which he stated was "possibly from bird ?Kuszholia sp. or from a theropod or a segnosaur", and this was identified as a therizinosauroid by Sues and Averianov (2016). He also figured proximal femur CCMGE 479/12457 as a theropod before it was referred to Caenagnathasia by Sues and Averianov (2015).
Funston et al. (2020b) report "a specimen of Caenagnathasia martinsoni that bore teeth was allegedly collected (J. Stiegler pers. comm.), but was lost in transit", which would be interesting as Funston et al. (2020a) concluded "the occlusal structures in caenagnathids cannot be homologous to alveoli, because they fail Patterson’s (1982) test of similarity by differing in developmental and histological nature."
Relationships- The holotype was first incorrectly listed as ZIN PO 3486 and identified as an ichthyornithid (Nessov, 1990). Nessov (1992) later named this as a new genus in its own family Kuszholiidae, under "suborder Theropoda + Aves." He viewed it as possibly a basal flightless bird like Patagopteryx or possibly a non-bird theropod which was convergent with birds. Nessov and Panteleev (1993) later assigned it to Patagopterygiformes, though their argument has yet to be translated from Russian. Kurochkin (2001) retained Kuszholia as Aves (sensu Chiappe) incertae sedis, and noted that patagopterygiform affinities were not yet verified. He did cite two supposed similarities though- enlarged third pair of sacral transverse processes and ventral sacrum convex. Yet Kuszholia's sacrum is ventrally concave (as in Patagopteryx and many other theropods) and the large transverse processes are on the second to last sacral. As all theropods have at least five sacrals, this corresponds to the fourth sacral or greater. All posterior sacral transverse processes are broken off in Patagopteryx in any case. Patagopteryx further differs in lacking sacral pleurocoels and having a convex posterior articular surface, as noted by Kurochkin. Kurochkin later (2006) placed Kuszholia in Euornithes (his Ornithurae), but outside Carinatae, in his phylogram, though without stated support.
The postzygapophyses of the last sacral are much larger than any avebrevicaudan, suggesting it is not a member of that clade. As noted above, dorsoventrally compressed centra are common in maniraptorans, while ventral grooves are present in the posterior sacrals of Ornitholestes and most maniraptoriforms. Very few maniraptorans have pleurocoels extending to the last sacral centrum, with examples limited to Neimongosaurus and caenagnathoids. Adult therizinosaurs are far larger, with even the smallest basal members (e.g. Beipiaosaurus) being four times as big, and the fusion does indicate Kuszholia's holotype is from an adult. There are small caenagnathoids though, including Caenagnathasia from the same formation. In fact, Caenagnathasia would have comparably sized sacrals to Kuszholia if scaled from other caenagnathoids. Note Sues and Averianov (2015) assigned the Kuszholia paratype and referred cervical to Caenagnathasia without commenting on why the holotype was also not referrable. Indeed, there seems to be no reason to refer any of the Bissekty caenagnathoid material to one genus instead of the other as the type synsacrum of Kuszholia cannot be compared to the type dentaries of Caenagnathasia. If they are synonymous, Kuszholia would have priority by two years, which is awkward as its synsacrum is less diagnostic and less phylogenetically determinable than Caenagnathasia's dentaries. The alternative is listing all material except the three dentaries and Kuszholia's holotype as Caenagnathoidea/idae indet., but tentative synonymization is chosen here.
The holotype is generally similar to Chirostenotes, but differs in having much larger pleurocoels and narrower postzygapophyses. It also has larger pleurocoels than Epichirostenotes, Citipes and Shixinggia. Avimimus is unique among oviraptorosaurs in lacking sacral pleurocoels, so is quite different. Funston and Currie (2016) found the composite OTU (not including the Kuszholia holotype) to be an elmisaurine caenagnathid between Caenagnathus and Chirostenotes. Adding taxa to Hartman et al.'s matrix recovers the Caenanagathasia dentaries in Avimimus, but the Kuszholia composite in Elmisaurinae as derived as Elmisaurus but outside Shixinggia+Avimimus.
References- Nessov, 1990. Small ichthyornithiform bird and other bird remains from Bissekty Formation (Upper Cretaceous) of central Kyzylkum Desert. Proceedings of the Zoological Institute, Leningrad. 210, 59-62.
Nessov, 1992. Review of localities and remains of Mesozoic and Paleogene birds of the USSR and the description of new findings. Russkii Ornitologicheskii Zhurnal. 1(1), 7-50.
Nessov and Panteleev, 1993. On the similarity of the Late Cretaceous ornithofauna of South America and Central Asia. Trudy Zoologicheskogo Instituta, RAN. 252, 84-94.
Currie, Godfrey and Nessov, 1993 (published 1994). New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30(10), 2255-2272.
Kurochkin, 1995. Synopsis of Mesozoic birds and early evolution of class Aves. Archaeopteryx. 13, 47-66.
Nessov, 1995. Dinosaurs of nothern Eurasia: New data about assemblages, ecology, and paleobiogeography. Institute for Scientific Research on the Earth's Crust, St. Petersburg State University, St. Petersburg. 1-156.
Nessov, 1997. Cretaceous nonmarine vertebrates of Northern Eurasia. Izdatelstvo Sankt-Peterburgskogo Universiteta, Saint Petersburg. 218 pp.
Kurochkin, 2001. Mesozoic birds of Mongolia and the former USSR. In Benton, Shishkin, Unwin and Kurochkin (eds.). The Age of Dinosaurs in Russia and Mongolia. 533-559.
Kurochkin, 2006. Parallel evolution of theropod dinosaurs and birds. Entomological Review. 86(suppl. 1), S45-S58.
Zelenkov and Averianov, 2011. Synsacrum of a primitive bird from the Upper Cretaceous of Uzbekistan. Paleontological Journal. 45(3), 314-319.
Sues and Averianov, 2015 (online 2014). New material of Caenagnathasia martinsoni (Dinosauria: Theropoda: Oviraptorosauria) from the Bissekty Formation (Upper Cretaceous: Turonian) of Uzbekistan. Cretaceous Research. 54, 50-59.
Sues and Averianov, 2016 (online 2015). Therizinosauroidea (Dinosauria: Theropoda) from the Upper Cretaceous of Uzbekistan. Cretaceous Research. 59, 155-178.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.
Funston, Wilkinson, Simon, Leblanc, Wosik and Currie, 2020a (online 2019). Histology of caenagnathid (Theropoda, Oviraptorosauria) dentaries and implications for development, ontogenetic edentulism, and taxonomy. The Anatomical Record. 303(4), 918-934.
Funston, Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020b. A new two-fingered dinosaur sheds light on the radiation of Oviraptorosauria. Royal Society Open Science. 7: 201184.
K. sp. (Yao, Wang, Sullivan, Wang, Stidham and Xu, 2015)
Middle-Late Campanian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China
Material- (IVPP V20377) anterior dentaries (symph 8.0 mm)
Comments- This was discovered in 2012 at "a rare microvertebrate locality within the Iren Dabasu Formation, about 16 km northeast of Erenhot City", which would put it in localities Q-T of Xing et al. (2012). Yao et al. (2015) referred IVPP V20377 to Caenagnathasia sp., and it does possess all characters here listed as diagnostic for that taxon. Of their characters listed as varying between Caenagnathasia specimens, most also vary between Chirostenotes specimens (posterior surface of symphysis with tubercle; chin-like eminence between anterior and ventral surfaces; pneumatic foramen in front of mandibular fenestra on lateral surface of dentary; depression on posteroventral margin of symphysis [ontogenetic?]), while paired second anterior occlusal grooves flanking first anterior occlusal groove is polymorphic in the holotype, and lateral projections on lingual ridges may be absent in ZIN PH 2354/16 due to preservation. This leaves presence of a median symphyseal groove on the posterodorsal depression in the holotype and IVPP V20377 but not ZIN PH 2354/16 (ontogenetic?) and on the posteroventral symphysis of IVPP V20377 but not the holotype or ZIN PH 2354/16 (taxonomic?). The age difference suggests the specimen is not conspecific with the Bissekty species.
References- Xing, He, Li and Xi, 2012. A review on the study of the stratigraphy, sedimentology, and paleontology of the Iren Dabasu Formation, Inner Mongolia. In Dong (ed.). Proceedings of the Thirteenth Annual Meeting of the Chinese Society of Vertebrate Paleontology. China Ocean Press. 1-44.
Yao, Wang, Sullivan, Wang, Stidham and Xu, 2015. Caenagnathasia sp. (Theropoda: Oviraptorosauria) from the Iren Dabasu Formation (Upper Cretaceous: Campanian) of Erenhot, Nei Mongol, China. Vertebrata PalAsiatica. 53(4), 291-298.
Wang, Zhang and Yang, 2018. Reevaluation of the dentary structures of caenagnathid oviraptorosaurs (Dinosauria, Theropoda). Scientific Reports. 8:391.

Caenagnathinae Sternberg, 1940 sensu Paul, 1988
Definition- (Caenagnathus collinsi <- Oviraptor philoceratops, Avimimus portentosus) (Martyniuk, 2012)
Other definitions- (Caenagnathus collinsi <- Elmisaurus elegans, Caenagnathasia martinsoni) (Longrich, Barnes, Clark and Millar, 2013)
(Caenagnathus collinsi <- Elmisaurus rarus) (Hendrickx, Hartman and Mateus, 2015)
Comments- Longrich et al. (2013) thought elegans was a caenagnathine, so no doubt meant to use Elmisaurus rarus as the external specifier instead.
References- Sternberg, 1940. A toothless bird from the Cretaceous of Alberta. Journal of Paleontology. 14(1), 81-85.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster: New York 464 pp.
Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs. Vernon, New Jersey. Pan Aves. 189 pp.
Longrich, Barnes, Clark and Millar, 2013. Caenagnathidae from the Upper Campanian Aguja Formation of west Texas, and a revision of the Caenagnathinae. Bulletin of the Peabody Museum of Natural History. 54(1), 23-49.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.

Anzu Lamanna, Sues, Schachner and Lyson, 2014
A. wyliei Lamanna, Sues, Schachner and Lyson, 2014
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US
Holotype- (CM 78000) (~3.5 m; ~284 kg) premaxillary fragments, quadrates, pterygoids, incomplete braincase, cranial fragments, incomplete mandibles (320 mm, symph 80 mm), six cervical vertebrae, dorsal ribs, gastralia, nine caudal vertebrae, twenty-eighth caudal vertebra (33.9 mm), twenty-ninth caudal vertebra (37.2 mm), partial thirtieth caudal vertebra, six chevrons, scapulocoracoids (scap 415 mm, cor 121 mm), humerus (345 mm), radius (280 mm), ulna (280 mm), phalanx I-1 (123.5 mm), manual ungual I (89.1 mm, 130 mm on curve), metacarpal II (139 mm), phalanx II-1 (131.8 mm), phalanx III-3 (82.8 mm), femora (525 mm), tibiae (660 mm), fibulae (585, 580 mm), astragalocalcanea (98.5, 98.0 mm transversely), metatarsals I (76 mm), phalanx I-1 (101.1 mm), phalanx III-1 (114.2 mm), pedal ungual III (101.1 mm), phalanx IV-2 (63.7 mm), phalanx IV-3 (52.5 mm), phalanx IV-4 (56.5 mm), five phalanges, two pedal unguals, partial metatarsal V
Paratypes- (CM 78001) premaxillae (one incomplete, one fragmentary), maxillae, jugals, quadrates, ectopterygoid, pterygoids, braincase, cranial fragments, atlas, axis, third cervical vertebra, fourth cervical vertebra, fifth cervical vertebra, sixth cervical vertebra, seventh cervical vertebra, eighth cervical vertebra, ninth cervical vertebra (~82.9 mm), tenth cervical vertebra, eleventh cervical vertebra (~78.5 mm), twelfth cervical vertebra, ten dorsal vertebrae (anterior 47 mm), seventeen ribs, eleven gastralia, incomplete sacrum (315 mm), twelve caudal vertebrae, eight chevrons, sternal plates (194.2 mm), ilia, pubes (450, 465 mm), ischia (305 mm), femora (505, 500 mm), tibiae (595 mm), fibulae (570 mm), astragalocalcanea (109.7 mm), phalanx III-3 (85.9 mm), pedal phalanx, four pedal unguals, metatarsal V (114.4 mm)
(FMNH PR2296 = BHM 2033) posterior mandible
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US
Referred- (BMRP 2013.4.1) dorsal ribs, caudal vertebra, pelvic elements, distal hindlimb including incomplete metatarsi (Holtz, Williams, Tremaine and Matthews, 2014)
?(MOR 9722) (~1.5-2 m) incomplete axial series, ribs, gastralia, partial forelimb including manual phalanges and unguals, ilia, partial pubes, ischia, partial tibia, tarsals, proximal metatarsals, pedal phalanges and pedal unguals (Flora, Wilson, Gardner and Fowler, 2015)
(ROM 65884) (8 year old subadult) three dorsal vertebral fragments, several dorsal rib fragments, several gastralial fragments, incomplete ~twenty-seventh caudal vertebrae (30.9 mm), ~twenty-eighth caudal vertebra (31.4 mm), ~twenty-ninth caudal vertebra (35.2 mm), distal manual phalanx II-1, pubic fragments, tibial fragments, incomplete fibula, incomplete pes (sold), partial metatarsal II, partial metatarsal III, partial metatarsal IV, metatarsal V, metatarsal fragments (Benner, Cullen and Evans, 2016)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, North Dakota, US
Paratype- (MRF 318) ninth cervical vertebra, eleventh cervical vertebra, twelfth cervical vertebra, dorsal rib, scapulocoracoid, radius (275 mm), ulna (280 mm)
Late Cretaceous?
USA?
Referred- (BDM uncatalogued) metatarsal I (Yun and Funston, 2021)
Diagnosis- (after Lamanna et al., 2014) tall, crescentic cranial crest formed by posterodorsal processes of premaxillae (unknown in other caenagnathids); body of maxilla lacking antorbital fossa (unknown in other caenagnathids except Epichirostenotes); maxillary ascending process elongate and shaped like an inverted L (unknown in other caenagnathids); quadratojugal process of jugal dorsoventrally deep (unknown in other caenagnathids); quadratojugal process of jugal bifurcated posteriorly (unknown in other caenagnathids); occipital condyle transversely wider than foramen magnum (unknown in other caenagnathids except Epichirostenotes); prominent lateral flange on symphyseal region of dentary (unknown in other caenagnathids except Caenagnathus, Chirostenotes, Elmisaurus, Citipes and Kuszholia); elongate retroarticular process of mandible (subequal in anteroposterior length to quadrate articulation) (unknown in other caenagnathids except for Caenagnathus, Chirostenotes and Apatoraptor) ; distal end of radius divided into two rounded processes (unknown in other caenagnathids); sulcus on ventromedial aspect of manual phalanx II-1 (unknown in other caenagnathids except Hagryphus, Chirostenotes and Elmisaurus); tubercle on anterior surface of astragalus near base of ascending process (unknown in other caenagnathids except Chirostenotes).
(after Funston and Currie, 2016) rounded hypapophyses in anterior dorsal vertebrae (unknown in other caenagnathids except Apatoraptor and Epichirostenotes); fenestra in posterolateral sternum (unknown in other caenagnathids except Apatoraptor).
Comments- Currie et al. (1993) described a posterior mandible (their BHM 2033) which was larger than any Dinosaur Park Caenagnathus and differed from C. collinsi and C. sternbergi in glenoid morphology. They referred it to Caenagnathus sp.. Varricchio (2001) further noted it shared several characters with C. collinsi to the exclusion of C. sternbergi, and Lamanna et al. (2014) referred it to the present species.
Triebold et al. (2000) reported two new large oviraptorosaur specimens discovered in 1998, identifying them as oviraptorids, though they were quickly reidentified as caenagnathid. These specimens were officially described by Lamanna et al. (2014) as the new taxon Anzu wyliei. The mandible is most similar to the Caenagnathus collinsi holotype.
Schachner et al. (2006; 2007) announced MRF 318 as Chirostenotes sp., which was later described as a specimen of Anzu by Lamanna et al. (2011; 2014).
Holtz et al. (2014, 2015) announced a new specimen (BMRP 2013.4.1- Holtz pers. comm, 2015) found in June 2013, which has only distal tarsal IV fused to the metatarsus, unfused metatarsals, and "a pair of pronounced cruciate ridges on the plantar surface of metatarsal III" which don't extend proximally as far as Elmisaurus.
ROM 65884 was found in 2012, and unfortunately a "nearly complete right foot was also collected, assembled into a display mount, and sold to a private collector" before the ROM bought the rest of the material (Cullen et al., 2020). Initially reported in an SVP abstract by Benner et al. (2016) as "a large-bodied taxon similar in size to the recently described, coeval taxon Anzu wyliei", Cullen et al. described the specimen as cf. Anzu wyliei, as it is from the same formation and almost identical in anatomy and size ("c. 3% difference in linear size, on average"). Indeed, the slightly curved metatarsal V is shared with Anzu but not Chirostenotes, Elmisaurus or seemingly Macrophalangia (+ Caenagnathus?). While this specimen was found in the same state as BMRP 2013.4.1 and preserves almost the same elements, they were discovered in different years and different counties and both metatarsi of BMRP 2013.4.1 were recovered by the Burpee Museum.
Yun and Funston (2021) figure the metatarsal I of the holotype in detail, as well as an additional Anzu metatarsal I called "BDM uncatalogued specimen," for which no other information is available in the literature.
Funston and Currie (2016) recover Anzu as basal to the caenagnathine-elmisaurine split, sister to Epichirostenotes.
References- Currie, Godfrey and Nessov, 1993 (published 1994). New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30(10), 2255-2272.
Triebold, Nuss and Nuss, 2000. Initial report of a new North American Oviraptor. In: The Florida Symposium on Dinosaur Bird Evolution, Presented by the Florida Institute of Paleontology at the Graves Museum of Archaeology and Natural History, Dania Beach, Florida, USA. p. 25.
Varricchio, 2001. Late Cretaceous oviraptorosaur (Theropoda) dinosaurs from Montana. In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life. New Research Inspired by the Paleontology of Philip J. Currie. Indiana University Press. 42-57.
Schachner, Lyson and Hanks, 2006. A preliminary report of a new specimen of Chirostenotes (Oviraptorosauria: Theropoda) from the Hell Creek Formation of North Dakota. Journal of Vertebrate Paleontology. 26, 120A.
Schachner, Lyson, Atterholt and Hanks, 2007. A preliminary report of a new specimen of Chirostenotes (Oviraptorosauria: Theropoda) from the Hell Creek Formation of North Dakota. Journal of Vertebrate Paleontology. 27(3), 141A.
Lamanna, Sues, Schachner and Lyson, 2011. A new caenagnathid oviraptorosaur (Theropoda: Maniraptora) from the Upper Cretaceous (Maastrichtian) Hell Creek Formation of the western United States. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 140.
Holtz, Williams, Tremaine and Matthews, 2014. New additions to the Hell Creek Formation (Upper Maastrichtian) vertebrate fauna of Carter County, Montana. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 149.
Lamanna, Sues, Schachner and Lyson, 2014. A new large-bodied oviraptorosaurian theropod dinosaur from the Latest Cretaceous of western North America. PLoS ONE. 9(3), e92022.
Flora, Wilson, Gardner and Fowler, 2015. A three-dimensionally articulated probable oviraptorosaur from the Hell Creek Formation of Montana. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 124-125.
Holtz, Williams and Tremaine, 2015. A new specimen of Anzu (Caenagnathidae, Oviraptorosauria): Implications for the proposed Caenagnathinae/Elmisaurinae division and for cursoriality in caenagnathids. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 146.
Benner, Cullen and Evans, 2016. A new large-bodied caenagnathid specimen (Theropoda, Oviraptorosauria) from the Hell Creek Formation (Late Cretaceous) of Montana, with implications for osteohistological variability in caenagnathids. Journal of Vertebrate Paleontology. Program and Abstracts, 96.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.
Cullen, Simon, Benner and Evans, online 2020. Morphology and osteohistology of a large-bodied caenagnathid (Theropoda: Oviraptorosauria) from the Hell Creek Formation (Montana): Implications for size-based classifications and growth reconstruction in theropods. Papers in Palaeontology. Early View. DOI: 10.1002/spp2.1302
Yun and Funston, 2021. A caenagnathid oviraptorosaur metatarsal from the Mesaverde Formation (Campanian) of Wyoming, USA. Vertebrate Anatomy Morphology Palaeontology. 9, 105-115.

Caenagnathus Sternberg, 1940
?= Macrophalangia Sternberg, 1932
C. collinsi Sternberg, 1940
?= Macrophalangia canadensis Sternberg, 1932
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada
Holotype- (CMN 8776) incomplete mandibles (230 mm, symph 61.9 mm) (femur ~377 mm)
Referred- ?(CMN 8538; holotype of Macrophalangia canadensis) distal tibia, partial astragalus, distal tarsal III, distal tarsal IV, metatarsal I (~47 mm), phalanx I-1 (58 mm), pedal ungual I (31 mm), metatarsal II (205 mm; lost), phalanx II-1 (78 mm), phalanx II-2 (63 mm), pedal ungual II (60 mm), partial metatarsal III (230 mm; lost), phalanx III-1 (75 mm), phalanx III-2 (52 mm), phalanx III-3 (58 mm), pedal ungual III (60 mm), metatarsal IV (212 mm; lost), phalanx IV-1 (59 mm), phalanx IV-2 (33 mm), phalanx IV-3 (31 mm), phalanx IV-4 (35 mm), pedal ungual IV, metatarsal V (60 mm) (Sternberg, 1932)
(CMN 12372; = CMN 12322 of Longrich et al.) manual ungual II (Longrich et al., 2013)
(TMP 1979.014.0001) manual ungual I (~90 mm on curve) (femur ~364 mm) (Funston, Persons, Bradley and Currie, 2015)
(TMP 1979.015.0001) manual ungual I (~69 mm) (Currie, 1992)
(TMP 1982.019.0222) manual ungual I (91 mm on curve) (femur ~369 mm) (Funston, Persons, Bradley and Currie, 2015)
?(TMP 1986.036.0323) (~96 kg) femur (370 mm) (Funston, Persons, Bradley and Currie, 2015)
?(TMP 1993.036.0197) (femur ~438 mm) metatarsal II (261 mm) (Funston, Persons, Bradley and Currie, 2015)
?(TMP 1993.036.0198) (femur ~411 mm) metatarsal II (245 mm) (Funston, Persons, Bradley and Currie, 2015)
(TMP 1993.0360.475) manual ungual II (108 mm on curve) (femur ~403 mm) (Bell, Currie and Russell, 2015)
(TMP 2009.003.0029; = TMP 1982.019.0222 in part) manual ungual I (85 mm on curve) (femur ~343 mm) (Bell, Currie and Russell, 2015)
(UALVP 55725) partial caudal vertebra (39.7 mm) (femur ~393 mm) (Funston, Persons, Bradley and Currie, 2015)
(UALVP 56638) (12 year old adult) pubes (416 mm) (Funston, 2020)
(UALVP 59791) partial ilium (Funston, 2020)
?(UALVP 59921) manual phalanx I-1 (102 mm) (Funston, 2020)
Diagnosis- (after Sternberg, 1940) elongate dentary symphysis (also in Anzu; unknown in other caenagnathids except Chirostenotes, Elmisaurus and Kuszholia).
(after Currie et al., 1993) dorsal midline ridge on anterior portion of mandibular symphysis (unknown in other caenagnathids except Chirostenotes, Elmisaurus and Kuszholia); lingual ridges converge to meet (unknown in other caenagnathids except Anzu, Chirostenotes, Elmisaurus, Citipes and Kuszholia); lateral occlusal grooves confluent with anterior occlusal grooves (also in Anzu; unknown in other caenagnathids except Chirostenotes, Elmisaurus and Kuszholia); lower articular ridge on mandible (also in Anzu and Apatoraptor; unknown in other caenagnathids except Chirostenotes); medial glenoid longer anteroposteriorly (also in Anzu and Apatoraptor; unknown in other caenagnathids except Chirostenotes).
(after Funston et al., 2015) slight concavity between proximal articular surface and flexor tubercle in manual ungual I (also in Anzu; unknown in other caenagnathids except Hagryphus, Chirostenotes and Elmisaurus); short and deep groove between proximal articular surface and flexor tubercle in manual ungual II (unknown in other caenagnathids except Chirostenotes and Elmisaurus); distally abrupt large posterior protuberance on proximal end of metatarsal II (unknown in other caenagnathids except Chirostenotes, Elmisaurus and Citipes); facet for metatarsal III on metatarsal II extends only about halfway up element (also in Citipes; unknown in other caenagnathids except Chirostenotes).
(after Funston and Currie, 2016) dentary-angular bar below external mandibular fenestra not bowed ventrally (unknown in other caenagnathids except Anzu, Chirostenotes and Apatoraptor).
(after Funston, 2020) low ilium above acetabulum (unknown in other caenagnathids except Chirostenotes and Citipes); rounded ventral edge of preacetabular blade (unknown in other caenagnathids except Anzu and Apatoraptor); inclined ventral edge of pubic peduncle of ilium (also in Chirostenotes; unknown in other caenagnathids except Anzu? and Citipes); pubic peduncle of ilium with anteroposterior ridge on ventral surface for contact with pubis, forming concavo-convex contact (unknown in other caenagnathids except Chirostenotes); pubes relatively straight in anterior view, producing transversely narrow proximal end (also in Epichirostenotes; unknown in other caenagnathids except Chirostenotes/Citipes TMP 1994.012.0603); iliac contact of pubis with anteroposterior concavity for ridge on the pubic peduncle of the ilium, forming concavo-convex contact (unknown in other caenagnathids except Chirostenotes/Citipes TMP 1994.012.0603).
(proposed) dentary posterodorsal process extends posteriorly past coronoid process (unknown in other caenagnathids except Anzu, Chirostenotes, Apatoraptor and Kuszholia); pedal phalanx III-3 longer than III-2 (unknown in other caenagnathids except Elmisaurus and Citipes).
Other diagnoses- Being the first described caenagnathid mandible and without relationships with Oviraptor being recognized for 36 more years, Sternberg's (1940) initial diagnosis included mostly characters plesiomorphic for caenagnathoids (no teeth; symphysis relatively broad; mandibular rami parallel; external mandibular fenestra large; articular surface large, oval and convex, with a ridged axis parallel to the direction of the ramus) and caenagnathids (dentaries fused at symphysis). The final character (retroarticular, and lateral and medial articular processes short) is relative to Aves and true of most Mesozoic theropods.
Funston et al. (2015) also listed "the anterior projection of the facet for metatarsal IV on the proximal end" of metatarsal II as being different from Chirostenotes, but this is comparable to TMP 1979.020.001.
Comments- Caenagnathus collinsi was named for a pair of mandibles found in Summer 1936, and assigned to Aves (Sternberg, 1940). This assignment was rejected by most paleornithologists such as Wetmore (1960), who hypothesized a relationship with ornithomimids. However, Cracraft (1971) supported an avian relationship, specifically with Galloanseres. In the same paper he named a new species of Caenagnathus, C. sternbergi, known from a posterior mandible. This differs from C. collinsi in a few characters, suggesting early that two species were present in the Dinosaur Park Formation. Currie et al. (1993) later described five dentaries which also differ from C. collinsi. They referred to these as Caenagnathus cf. sternbergi, as none were directly comparable to the C. sternbergi holotype. A mandible discovered in 2001 (TMP 2001.012.0012) and described by Funston and Currie (2014) eventually verified Currie et al.'s dentaries went with sternbergi articulars. Caenagnathus' identity was finally solved by Osmólska (1976), who allied it with oviraptorids. Currie and Russell (1988) first suggested Caenagnathus and Chirostenotes (then only known from postcrania) were synonymous, which was strengthened by Sues' (1994) Horseshoe Canyon specimen with Chirostenotes-like postcrania and an edentulous maxilla and shown to be true at the family level by Anzu (Lamanna et al., 2011) which preserved skeletons with Caenagnathus-like mandibles. Longrich et al. (2013) separated Caenagnathus collinsi from Chirostenotes pergracilis based on differences between the type mandible of the former and sternbergi, which they synonymize with pergracilis (as partial skeleton UALVP 59400 later confirmed). They referred manual ungual CMN 12372 ("erroneously reported as CMN 12322 by those authors" as stated by Bell et al., 2015) based on "its large size and robust construction." Bell et al. (2015) described two additional manual unguals (TMP 1993.036.0475 and 2009.003.0029), noting Caenagnathus "based on size alone poses a conceivable match for the Dinosaur Park unguals described here." Funston et al. (2015) agreed Caenagnathus collinsi is distinct from Chirostenotes pergracilis, describing several new specimens as Caenagnathus (TMP 79.14.1, 82.19.222, 86.36.323, 93.36.197, 93.36.198, 93.36.631, 93.75.49 and UALVP 55725). Funston (2020) later considered referral of femora (especially TMP 1986.036.0323) to be more ambiguous due to his histological analyses of Chirostenotes indicating it may have grown larger than previously expected. Funston also redescribed the holotype mandible and referred a large ilium (UALVP 59791) and pubis (UALVP 56638), the former of which is lower than Chirostenotes or Citipes and has a derived pubic contact not present in Chirostenotes. The ilium and pubis are described in detail and figured in Funston's (2019) thesis, where he notes regarding UALVP 56638 "no locality data accompanies the specimen because it was mislabelled, but the excellent preservation and associated matrix are consistent with the DPF." Of the new UALVP specimens, 55725 was found on May 31 2014, 59791 May 27 2018 and 59921 July 8 2019 (UALVP online).
Sternberg (1932) described Macrophalangia canadensis based on a pes found in 1928, then thought to be an ornithomimid. Colbert and Russell (1969) noted it may be synonymous with Chirostenotes, though this was not shown to be likely until Osmólska (1981) described Elmisaurus which preserved a manus and pes of similar morphology. Longrich et al. (2013) proposed Macrophalangia was synonymous with Caenagnathus collinsi based on size and robusticity compared to Chirostenotes pergracilis, which they viewed as a separate taxon. Similar or greater differences in size and robusticity are known for other coelurosaur species so this is not very convincing evidence. Funston (2020) referred Macrophalangia to Chirostenotes, but noted it differed from TMP 1979.020.0001 in having a less constricted metatarsal I (waist 95% of proximal depth vs. 76%), pedal phalanx I-1 longer than metatarsal I (123% vs. 94%) and a straight metatarsal V (unillustrated in Macrophalangia but described as a "small round splint"). The former two characters were said to be shared with Anzu, but while phalanx I-1 length (133%) is even greater than Macrophalangia, metatarsal I constriction (82%) and metatarsal V curvature are both intermediate in Anzu. Funston (pers. comm. 2015) noted the metatarsals of Macrophalangia are lost, having gone on loan and never being returned. It is here provisionally referred to Caenagnathus but as in Longrich et al. is not officially considered a senior synonym pending more information (e.g. histology of CMN 8538).
Currie (1992) reported "an isolated ungual from the Judith River Formation of Alberta [that] is unusual in that the flexor tubercle is well developed and extends posteriorly below the articulation with the penultimate phalanx. The only Cretaceous animal that I am aware of with this unusual articulation is Segnosaurus (Perle, 1981)." This last reference is actually for Erlikosaurus, whose pedal unguals match this description, while the manual unguals of Segnosaurus are unfigured and its preserved pedal ungual is only figured in dorsal view. TMP 1979.015.0001 was figured as "possible segnosaurid ungual." Cullen et al. (2020) found this specimen "to be most similar in morphology to caenagnathids", which was predicted by Currie in the initial 1992 publication, where he wrote "the ungual resembles the manual ungual of Chirostenotes if the articulation is ignored, and it is possible that the distinctive nature of the articulation may have been produced by disease or injury." Given the presence of three Dinosaur Park caenagnathids, I refer TMP 1979.015.0001 to Caenagnathus based on its large size.
References- Sternberg, 1932. Two new theropod dinosaurs from the Belly River Formation of Alberta. Canadian Field-Naturalist. 46(5), 99-105.
Sternberg, 1940. A toothless bird from the Cretaceous of Alberta. Journal of Paleontology. 14(1), 81-85.
Wetmore, 1960. A classification for the birds of the world. Smithsonian Miscellaneous Collections. 139(11), 1-37.
Colbert and Russell, 1969. The small Cretaceous dinosaur Dromaeosaurus. American Museum Novitiates. 2380, 1-49.
Cracraft, 1971. Caenagnathiformes: Cretaceous birds convergent in jaw mechanism to dicynodont reptiles. Journal of Paleontology. 45(5), 805-809.
Osmólska, 1976. New light on skull anatomy and systematic position of Oviraptor. Nature. 262, 683-684.
Osmólska, 1981. Coossified tarsometatarsi in theropod dinosaurs and their bearing on the problem of bird origins. Palaeontologia Polonica. 42, 79-95.
Currie and Russell, 1988. Osteology and relationships of Chirostenotes pergracilis (Saurischia, Theropoda) from the Judith River (Oldman) Formation of Alberta, Canada. Canadian Journal of Earth Sciences. 25(7), 972-986.
Currie, 1992. Saurischian dinosaurs of the Late Cretaceous of Asia and North America. In Mateer and Chen (eds.). Aspects of Nonmarine Cretaceous Geology. China Ocean Press. 237-249.
Currie, Godfrey and Nessov, 1993 (published 1994). New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30(10), 2255-2272.
Sues, 1994. New evidence concerning the phylogenetic position of Chirostenotes (Dinosauria: Theropoda). Journal of Vertebrate Paleontology. 14(3), 48A.
Lamanna, Sues, Schachner and Lyson, 2011. A new caenagnathid oviraptorosaur (Theropoda: Maniraptora) from the Upper Cretaceous (Maastrichtian) Hell Creek Formation of the western United States. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 140.
Longrich, Barnes, Clark and Millar, 2013. Caenagnathidae from the Upper Campanian Aguja Formation of west Texas, and a revision of the Caenagnathinae. Bulletin of the Peabody Museum of Natural History. 54(1), 23-49.
Funston and Currie, 2014. A previously undescribed caenagnathid mandible from the late Campanian of Alberta, and insights into the diet of Chirostenotes pergracilis (Dinosauria: Oviraptorosauria). Canadian Journal of Earth Sciences. 51(2), 156-165.
Bell, Currie and Russell, 2015 (online 2014). Large caenagnathids (Dinosauria, Oviraptorosauria) from the uppermost Cretaceous of western Canada. Cretaceous Research. 52, 101-107.
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of Alberta, Canada. Cretaceous Research. 54, 179-187.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria (Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Cullen, Larson, Zanno, Currie and Evans, 2020. Theropod biodiversity patterns in the Dinosaur Park Formation (Late Cretaceous: Campanian) of Alberta revealed through morphometrics and biostratigraphy. The Society of Vertebrate Paleontology 80th Annual Meeting, Conference Program. 115.
Funston, 2020. Caenagnathids of the Dinosaur Park Formation (Campanian) of Alberta, Canada: Anatomy, osteohistology, taxonomy, and evolution. Vertebrate Anatomy Morphology Palaeontology. 8, 105-153.
Rhodes, Funston and Currie, 2020. New material reveals the pelvic morphology of Caenagnathidae (Theropoda, Oviraptorosauria). Cretaceous Research. 114, 104521.

Epichirostenotes Sullivan, Jasinski and van Tomme, 2011
E. curriei Sullivan, Jasinski and van Tomme, 2011
Early Maastrichtian, Late Cretaceous
Horsethief Member of Horseshoe Canyon Formation, Alberta, Canada
Holotype- (ROM 43250) (adult) maxilla, palatine, braincase, third cervical vertebra, sixth cervical vertebra (85 mm), seventh cervical vertebra (81 mm), two cervical ribs, anterior dorsal vertebra (40 mm), anterior dorsal vertebra, dorsal rib, gastralia fragments, sacrum (270 mm), proximal caudal vertebra, four distal caudal vertebrae, ilial fragments, pubes (422 mm), ischium (212 mm), distal tibia
Middle Maastrichtian, Late Cretaceous
Tolman Member of Horseshoe Canyon Formation, Alberta, Canada
Referred- ?(CMN 9570) metatarsal II (258 mm) (Russell, 1984)
Diagnosis- (after Sullivan et al., 2011) ischium tapers gradually after obturator process (also in Anzu and Elmisaurus; unknown in other caenagnathids except Chirostenotes); ventral edge of ischium distal to obturator process shallowly concave (also in Anzu; unknown in other caenagnathids except Chirostenotes and Elmisaurus); triangular obturator process (distal edge <20% of obturator depth) (also in Anzu and Elmisaurus; unknown in other caenagnathids except Chirostenotes); ventrodistally curved pubic peduncle on ischium (also in Anzu; unknown in other caenagnathids except Chirostenotes).
(proposed) presence of an antorbital fossa (unknown in other caenagnathids except Anzu); larger foramen magnum than occipital condyle (unknown in other caenagnathids except Anzu); ventrally directed paroccipital processes (unknown in other caenagnathids except Anzu).
Other diagnoses- Sullivan et al. (2011) also listed "braincase much deeper than long, with distinctly verticalized basicranial region" which seems to be shared with Anzu and unknown in other caenagnathids. Similarly, "otic region with deep, but anteroposteriorly narrow, lateral depression" has not been evalauated in Anzu, and is not preserved in other taxa. "Synsacrum composed of six co-ossified vertebrae and pneumatic foramina" is also true in Anzu and Chirostenotes in regard to sacral number, and these two taxa, Citipes and Kuszholia in regard to all sacrals being pleurocoelous.
Comments- ROM 43250 was discovered on June 12 1923 and identified as an ornithomimid. Russell (1972) listed it as an undetermined ornithomimid, though Sues (1994) correctly identified it as caenagnathid in an SVP abstract before describing it in detail (Sues, 1997). Sullivan et al. (2011) proposed the name Epichirostenotes curriei for ROM 43250, based on stratigraphy and several ischial differences from TMP 1979.020.0001. These ischial characters may prove to be diagnostic (in particular, the more abrupt tapering is present in two Chirostenotes specimens), but they are comparable to interspecific variation in Tyrannosaurus rex specimens (AMNH 5027, CM 9380 and FMNH PR2081) so could prove to be questionable as more specimens are discovered. It is largely incomparable to the similarly sized Caenagnathus, although the pubic shaft is much more slender in side view. This may be due to the extensive taphonomic crushing described by Sues (1997) however.
CMN 9570 was discovered in 1926 and "was referred to Macrophalangia in the records of the National Museum of Natural Sciences (Ottawa) by Wann Langston, Jr." before being described as cf. Macrophalangia by Russell (1984). Currie and Russell (1988) illustrated it and referred the element to Chirostenotes sp.. Two caenagnathid taxa have since been identified from the Horseshoe Canyon Formation- Apatoraptor and Epichirostenotes, both based on partial skeletons lacking metatarsi. Funston (2019) tentaively refers it to the latter based on size.
References- Russell, 1972. Ostrich dinosaurs from the Late Cretaceous of western Canada. Canadian Journal of Earth Sciences. 9, 375-402.
Russell, 1984. A check list of the families and genera of North American dinosaurs. Syllogeus. 53, 1-35.
Currie and Russell, 1988. Osteology and relationships of Chirostenotes pergracilis (Saurischia, Theropoda) from the Judith River (Oldman) Formation of Alberta, Canada. Canadian Journal of Earth Sciences. 25(7), 972-986.
Sues, 1994. New evidence concerning the phylogenetic position of Chirostenotes (Dinosauria: Theropoda): Journal of Vertebrate Paleontology. 14(3), 48A.
Sues, 1997. On Chirostenotes, a Late Cretaceous oviraptorosaur (Dinosauria: Theropoda) from western North America. Journal of Vertebrate Paleontology. 17(4), 698-716.
Sullivan, Jasinski and van Tomme, 2011. A new caenagnathid Ojoraptorsaurus boerei, n. gen., n. sp. (Dinosauria, Oviraptorosauria), from the Upper Cretaceous Ojo Alamo Formation (Naashoibito Member), San Juan Basin, New Mexico. New Mexico Museum of Natural History and Science Bulletin. 53, 418-428.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria (Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.

Hagryphus Zanno and Sampson, 2005
H. giganteus Zanno and Sampson, 2005
Late Campanian, Late Cretaceous
Kaiparowitz Formation, Utah, US
Holotype- (UMNH VP 12765) distal radius, scapholunare, ulnare, semilunate carpal, distal carpal III, metacarpal I (66 mm), phalanx I-1 (87 mm), manual ungual I (94 mm on curve), metacarpal II (122 mm), phalanx II-1 (95 mm), phalanx II-2 (99 mm), metacarpal III (87 mm), phalanx III-1 (54 mm), phalanx III-2 (45 mm), phalanx III-3 (60 mm), manual ungual III (75 mm on curve), manual claw III impression, proximal pedal ungual I, distal metatarsal II, distal phalanx II-1, phalanx II-2, pedal ungual II, incomplete pedal ungual III, distal metatarsal IV, incomplete pedal ungual IV, fragmentary pedal phalanges
Referred- ?(RAM 12433) distal metatarsal II (Zanno, Loewen, Farke, Kim, Claessens and McGarrity, 2013)
Diagnosis- (after Zanno and Sampson, 2005) larger than Chirostenotes (also in Caenagnathus and Anzu); robust metacarpal I (transverse shaft width ~20% of length) (unknown in other caenagnathids except Chirostenotes, Apatoraptor and Elmisaurus); robust manual phalanx I-1 (transverse shaft width ~14% of length) (unknown in other caenagnathids except Anzu, Caenagnathus, Chirostenotes and Elmisaurus); interphalangeal joint of manual digit I extending just distal to the metacarpophalangeal joint of digit II (unknown in other caenagnathids except Chirostenotes, Apatoraptor and Elmisaurus).
Comments- The holotype was discovered in 2002 (Zanno et al., 2013). Being of similar size to the contemporaneous Caenagnathus collinsi, it differs in having a much more robust manual phalanx I-1 and a groove between the articular facet and flexor tubercle of manual ungual I.
Zanno and Sampson (2005) conservatively avoided the Chirostenotes/Caenagnathus versus Elmisaurus controversy and merely referred Hagryphus to Oviraptorosauria, though believed it to be most similar to those two genera. Funston and Currie (2016) recovered it more basal in Caenagnathidae than other Late Cretaceous American taxa, though closer than Microvenator and Gigantoraptor.
References- Zanno and Sampson, 2003. A new caenagnathid specimen from the Kaiprowits Formation (Late Campanian) of Utah. Journal of Vertebrate Paleontology. 23(3), 114A.
Zanno and Sampson, 2005. A new oviraptorosaur (Theropoda: Maniraptora) from the Late Cretaceous (Campanian) of Utah. Journal of Vertebrate Paleontology. 25(4), 897-904.
Zanno, Loewen, Farke, Kim, Claessens and McGarrity, 2013. Late Cretaceous theropod dinosaurs of southern Utah. In Titus and Loewen (eds.). At the Top of the Grand Staircase: The Late Cretaceous of Southern Utah. Indiana University Press. 504-525.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.

Apatoraptor Funston and Currie, 2016
A. pennatus Funston and Currie, 2016
Early Maastrichtian, Late Cretaceous
Horsethief Member of Horseshoe Canyon Formation, Alberta, Canada
Holotype- (TMP 1993.051.0001) (~80 kg, 2 year old subadult) palatine (~25 mm), incomplete mandible (~165 mm), hyoid (68.4 mm), (cervical series 680 mm), axial fragment, third to twelve cervical vertebrae fused to ribs (seventh 53.2, ninth 61.5 mm), nine dorsal vertebrae, nine dorsal ribs, five uncinate processes, scapulae (one proximal; 215 mm), coracoids, sternal plates (one fragmentary; 112 mm), sternal ribs, humerus (206 mm), radius, ulna (167 mm), (?)semilunate carpal, metacarpal I (52 mm), phalanx I-1 (82.4 mm), manual ungual I (50 mm on curve), metacarpal II (105 mm), phalanx II-1 (74 mm), phalanx II-2 (68.5 mm), manual ungual II (70 mm on curve), metacarpal III (75 mm), partial phalanx III-1, phalanx III-2 (32 mm), incomplete phalanx III-3, ilial fragment, partial femur (~345 mm), proximal tibia, proximal fibula
Diagnosis- (after Funston and Currie, 2016) anterior constriction of external mandibular fenestra by posteroventral dentary process (unknown in other caenagnathids except Anzu, Caenagnathus, Chirostenotes, Elmisaurus and Kuszholia); articular region of mandible with low articular ridge offset from dorsal margin of articular-surangular-coronoid complex (also in Anzu; unknown in other caenagnathids except Caenagnathus and Chirostenotes); small transverse processes on third cervical (also in Kuszholia?; unknown in other caenagnathids except Epichirostenotes); no postzygodiapophyseal lamina on sixth cervical (unknown in other caenagnathids except Epichirostenotes); parapophyses of sixth and seventh cervicals extend far anteriorly and are separated by a deep depression (unknown in other caenagnathids except Epichirostenotes); deep infraprezygapophyseal fossa in sixth and seventh cervicals? (also in Kuszholia?; unknown in other caenagnathids except Epichirostenotes); infradiapophyseal fossa present in fifth-eighth cervicals (unknown in other caenagnathids except Epichirostenotes); lamina connecting tuberculum and capitulum with corresponding pneumatic foramen in mid dorsal ribs (unknown in other caenagnathids except Chirostenotes?, Epichirostenotes and Elmisaurus); less curved posterior coracoid margin (also in Anzu; unknown in other caenagnathids except Chirostenotes); coracoid tubercle placed less than halfway down coracoid (unknown in other caenagnathids except Chirostenotes); metacarpal I less than half as long as metacarpal II (unknown in other caenagnathids except Hagryphus and Elmisaurus); manual phalanx I-1 longer than II-1 or II-2 (unknown in other caenagnathids except Hagryphus, Anzu, Chirostenotes and Elmisaurus); manual phalanx II-1 longer than II-2 (unknown in other caenagnathids except Hagryphus, Chirostenotes and Elmisaurus).
Other diagnoses- Differences between palatines in this and Epichirostenotes (longer anteroventral process, anterodorsal process less upturned) noted by Funston and Currie (2016) may be due to incompletely preserved edges in both. They stated "the dentary-surangular suture does not extend posteriorly to the coronoid process", supposedly unlike other caenagnathids, but this is true in Chirostenotes and Anzu as well. Funston and Currie also said that "the laminae connecting the neural spine to the transverse processes of C6 and C7 are not as well developed in Apatoraptor as in Epichirostenotes, especially in their posterior extent", but it is uncertain which laminae these would be since oviraptorosaurs lack cervical spinodiapophyseal laminae. Funston and Currie list "ventral flange of angular underlying posteroventral dentary process", but this seems true in other caenagnathids (Anzu, Chirostenotes TMP 20010.012.0012) with its seeming absence in the one preserved side of Caenagnathus being due to a dorsally displaced angular. They also list "medial fossa anterior to articular region of mandible" which is present in Caenagnathus, and unknown in other caenagnathids (unreported but possible to check in Chirostenotes and Anzu). The lack of rib fusion in the third cervical unlike Epichirostenotes (and Anzu) could be ontogenetic based on size.
Comments- Funston (2014) comments on TMP 1993.051.0001, a specimen discovered in 1993 and first believed to be ornithomimid. It is listed in Funston et al. (2015) as "unnamed taxon", is said to be from the Horseshoe Canyon Formation by Eberth et al. (2013) and tentatively assigned to Epichirostenotes there. Currie et al. (2016) incorrectly state it's from the Dinosaur Park Formation however, when commenting on its scapulocoracoid and humerus. The specimen was officially named and described by Funston and Currie (2016), recovered as closer to Elmisaurus rarus than to Citipes, Caenagnathasiaor any of the larger caenagnathids. Funston's (2019) description in his thesis includes CT scans not available in the published version. Simon and Evans (2021) via histological study found the specimen is a young subadult and proposed it may be a young Epichirostenotes, which is based on a larger specimen from the same formation. This would make the proposed cervical differences either ontogenetic or individual variation.
References- Eberth, Evans, Brinkman, Therrien, Tanke and Russell, 2013. Dinosaur biostratigraphy of the Edmonton Group (Upper Cretaceous), Alberta, Canada: Evidence for climate influence. Canadian Journal of Earth Sciences. 50(7), 701-726.
Funston, 2014. Understanding Alberta’s caenagnathids: Insights on anatomy and taxonomy from a new, articulated specimen. R.E. Peter Biology Conference. Abstract 10.
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of Alberta, Canada. Cretaceous Research. 54, 179-187.
Currie, Funston and Osmólska, 2016 (online 2015). New specimens of the crested theropod dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica. 61(1), 143-157.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria (Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Simon and Evans, 2021. Osteohistology supports immature ontogenetic status of North American oviraptorosaurs Apatoraptor pennatus and Chirostenotes pergracilis. The Society of Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual Meeting. 236-237.

Chirostenotes Gilmore, 1924
= "Steneodactylus" Gilmore vide Holtz, DML 1998
C. pergracilis Gilmore, 1924
= Caenagnathus sternbergi Cracraft, 1971
= "Steneodactylus pergracilis" Gilmore vide Holtz, DML 1998
= Chirostenotes sternbergi (Cracraft, 1971) Snively, Currie, Brinkman, Ryan, Braman, Gardner, Lam, Spivak and Neuman, 2001
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada
Holotype- (CMN 2367) distal metacarpal I, phalanges I-1 (one incomplete; 63 mm), manual unguals I (44 mm), partial metacarpal II, phalanges II-1 (one incomplete; 65 mm), phalanges II-2 (one incomplete; 72 mm), manual ungual II (62 mm), distal phalanges III-3 (44 mm; one lost), manual unguals III (36 mm; one lost)
Referred- (CMN 2690; holotype of Caenagnathus sternbergi) posterior mandible (Cracraft, 1971)
?(TMP 1979.014.0499) manual ungual II (83 mm) (Currie and Russell, 1988)
(TMP 1979.020.0001) (~67 kg, 2 year old juvenile) (sacrum- 200 mm) first sacral vertebra (36.9 mm), second sacral vertebra, third sacral vertebra, fourth sacral vertebra, fifth sacral vertebra, sixth sacral verebra (29.9 mm), proximal mid dorsal rib, three dorsal rib shafts, coracoid (58.1 mm tall), distal metacarpal I, phalanx I-1 (65.4 mm), manual ungual I (48 mm on curve), phalanx II-1 (71.9 mm), phalanx II-2 (75.8 mm), manual ungual II (69 mm; 83 mm on curve), phalanx III-1 (30.3 mm; lost), phalanx III-3 (39.3 mm), ilium (255 mm), ischium (138 mm), femur (304 mm), tibia (367 mm), metatarsal I (42.4 mm), phalanx I-1 (39.8 mm), distal tarsal III fragments, distal tarsal IV, metatarsal II (181 mm), metatarsal III (207 mm), phalanx III-1 (54.1 mm), metatarsal IV (186 mm), metatarsal V (33 mm) (Currie and Russell, 1988)
(TMP 1985.043.0070) (1 year old juvenile) anterior dentaries (symph 19 mm) (Funston, Wilkinson, Simon, Leblanc, Wosik and Currie, 2020)
(TMP 1990.056.0006) dentaries (symph 46.5 mm) (Currie, Godfrey and Nessov, 1993)
?(TMP 1991.144.0001) incomplete dentaries (symph 27.2 mm) (Currie, Godfrey and Nessov, 1993)
?(TMP 1992.036.0390) (juvenile?) incomplete dentaries (symph 29.6 mm) (Currie, Godfrey and Nessov, 1993)
(TMP 1992.036.1237) (7 year old subadult) anterior dentaries (symph 38.5 mm) (Funston, Wilkinson, Simon, Leblanc, Wosik and Currie, 2020)
?(TMP 1992.040.0044) (juvenile?) anterior dentaries (symph 32 mm) (Currie, Godfrey and Nessov, 1993)
(TMP 1993.036.0181) distal tarsal III, distal tarsal IV, metatarsal II (221 mm), metatarsal IV (221 mm) (Funston and Currie, 2014b; described by Funston et al., 2016)
(TMP 1996.012.0142) partial dentaries (symph 25.3 mm) (Funston, 2020)
(TMP 2001.012.0012) incomplete mandibles (188 mm; symph 32.8 mm) (Currie, 2005; described by Funston and Currie, 2014a)
(TMP 2002.012.0103) incomplete ilium (Funston and Currie, 2021)
(UALVP 59400) (9 year old adult) incomplete mandibles, seventh cervical vertebra (~49 mm), eighth cervical vertebra (~57 mm), fragmentary ninth cervical vertebra, cervical vertebra, fifth caudal vertebra, sixth caudal vertebra, seventh caudal vertebra, eighth caudal vertebra (27.1 mm), ninth caudal vertebra (25.2 mm), tenth caudal vertebra (22.3 mm), eleventh caudal vertebra (21.4 mm), twelfth caudal vertebra (19.5 mm), thirteenth caudal vertebra (19.4 mm), fourteenth caudal vertebra (16.1 mm), fifteenth caudal vertebra (20.4 mm), sixteenth caudal vertebra (20.9 mm), seventeenth caudal vertebra (16.9 mm), nine chevrons, proximal pubic fragment, fragmentary ischia, femoral and/or tibial fregments, distal tibia, partial astragalocalcanea, distal tarsal IV, feathers (Funston and Currie, 2021)
Middle-Late Campanian, Late Cretaceous
Belly River Group, Alberta, Canada
?(TMP 1979.008.0622) (juvenile?) anterior dentaries (symph 21 mm) (Currie, Godfrey and Nessov, 1993)
Late Campanian, Late Cretaceous
Upper Two Medicine Formation, Montana, US
?(MOR 1107) articular (symph ~28.5 mm scaled from TMP 2001.012.0012) (Varricchio, 2001)
Diagnosis- (after Cracraft, 1971) smaller than coeval Caenagnathus collinsi; articular ridge higher (unknown in other caenagnathids except Anzu, Caenagnathus and Apatoraptor); articular medial process not as elongated anteroposteriorly (unknown in other caenagnathids except Anzu, Caenagnathus and Apatoraptor).
(after Currie, 1989) metatarsal IV much shorter than III (<93%) (also in Caenagnathus?; unknown in other caenagnathids except Eoneophron, Elmisaurus and Citipes).
(after Varricchio, 2001) pedal phalanx III-1 >30% of metatarsal II in length (also in Caenagnathus?; unknown in other caenagnathids except Elmisaurus and Citipes).
(after Longrich et al., 2013) short dentary symphysis (median length <100% of transverse width at posterior edge), causing moderately divergent lateral dentary edges (~30-35 degrees) (also in Citipes, Elmisaurus and Kuszholia; unknown in other caenagnathids except Anzu and Caenagnathus); tip of beak with four lingual ridges (also in Elmisaurus and some Kuszholia; unknown in other caenagnathids except Caenagnathus; absent in TMP 1990.056.0006); dorsal edge of external mandibular fenestra everted laterally (unknown in other caenagnathids except Anzu, Caenagnathus and Apatoraptor); manual ungual I slender and weakly arched, flexor tubercles distally positioned (unknown in other caenagnathids except Hagryphus, Anzu, Caenagnathus, Apatoraptor and Elmisaurus).
(after Funston, 2020) occlusal tip of dentary upturned at approximately 45 degrees (taphonomically absent in TMP 1990.056.0006?; also in Elmisaurus and Kuszholia; unknown in other caenagnathids except Caenagnathus); posterior chevrons anteroposteriorly elongate at proximal end, as long or longer anteroposteriorly than corresponding caudal vertebrae (unknown in other caenagnathids except Anzu); tall ilium (height above acetabulum subequal to length between ventral ischial peduncle tip and anterior edge of pubic peduncle) (also in Citipes; unknown in other caenagnathids except Caenagnathus); distal tarsals and proximal metatarsals not coossified at maturity (also in Caenagnathus and metatarsals of Anzu; unknown in other caenagnathids except Eoneophron, Elmisaurus and Citipes); metatarsal V strongly procurving (unknown in other caenagnathids except Anzu, Ceanagnathus?, Elmisaurus and Citipes).
Other diagnoses- (after Gilmore, 1924) differs from Struthiomimus in shorter manual digits I and III, shorter metacarpal I, deeply ginglymoid metacarpals I and II, less medially divergent digit I, longer phalanx II-1, and more curved unguals. The shorter metacarpal I, and deeply ginglymoid metacarpals I and II also diagnosed to compared to Ornithomimus. The supposedly different deeply ginglymoid metacarpal III and short phalanges of digit III in Dromaeosaurus are due to mistaking its metatarsal II and pedal phalanges. It differs from Ornitholestes (including Tanycolagreus manus AMNH 587) in the longer metacarpal and phalanx I-1, and more divergent I-1.
Cracraft (1971) included the following character in his diagnosis compared to Caenagnathus collinsi's holotype, but this does not seem true of e.g. TMP 2001.012.0012- "portion of ramus immediately anterior to articular region much less robust and apparently relatively thinner dorsoventrally."
Almost all of Currie and Russell's (1988) characters are now known to be typical of caenagnathids, and often caenagnathoids in general- six sacral vertebrae with pleurocoels; digit III of manus longer than digit I, but with slender phalanges, having a diameter of less than half that of phalanges in other digits; well-developed posterodorsal lip on manual unguals; dolichoiliac, propubic pelvis; preacetabular portion of iliac blade longer than postacetabular; metatarsal III is proximally pinched between metatarsals II and IV, but only the proximal tip is excluded from the anterior surface of the metatarsus. Their final character, tarsometatarsus may have fused in old individuals, is currently considered false.
Currie (1989) proposed that in Citipes elegans the "preserved portion of metatarsal III is triangular in section, in contrast with Chirostenotes pergracilis, where it is diamond shaped", but these were being measured at different places along the shaft (see comments under Leptorhynchos). He also proposed a short metatarsal II differed from Elmisaurus rarus and elegans, but elegans specimens TMP 1982.016.0006 and TMP 2000.012.0008 have short metatarsals II as well (89% of mtIII) (as do Kol and Macrophalangia, the latter of which Currie viewed as synonymous with Chirostenotes).
Following the identification of dentaries used here, Longrich et al.'s 'Leptorhynchus elegans' examples are actually Chirostenotes pergracilis. As such, the anteriorly projecting beak tip in some specimens is individual or taphonomic variation, as is the varience of posterior divergence between dentaries (~30-35 degrees), and the symphyseal length/width ratio (49-87% vs. 58% in Citipes and 128% in Caenagnathus), although the divergence and symphyseal width are still different from Caenagnathus. Contrary to Longrich et al., the dentary is not excluded from the dorsal margin of the external mandibular fenestra by the surangular (TMP 2001.012.0012, 1990.056.0006), instead being similar to Caenagnathus with a small dorsal border. The beak tip can also be as elongate as Caenagnathus (e.g. UALVP 59400, TMP 1990.056.0006). Finally, the manual phalanges are not extremely long and slender compared to the one known specimen of Caenagnathus (I-1 UALVP 59921).
Varricchio (2001) listed chorda tympani foramen/slot absent as diagnostic of Caenagnathus sternbergi compared to C. collinsi, which Currie et al. (1993) had restoted CMN 8776 with. However, the feature is just an irregular transverse crack on each side at the posterior base of the glenoid (photo courtesy of McFeeters) instead of the anteroposterior groove on the dorsal glenoid surface which Currie et al. illustrated in Anzu specimen BHM 2033.
Funston (2020) included dentary fusion as a character, but this is true of all caenagnathids. A deep lateral dentary fossa is only present in TMP 2001.012.0012. Funston also listed cervical vertebrae with low neural spines and small epipophyses as diagnostic, but this is true for all oviraptorosaurs. Distal caudal vertebrae with anteriorly directed transverse processes are also present in Nomingia and Citipati, with the condition in Anzu (the only other caenagnathid to preserve the area) unreported.
Comments- Chirostenotes pergracilis was named in 1924 based on two manus collected in 1914, and thought by Gilmore to be a a coelurid intermediate between Ornitholestes and ornithomimids. He also referred a pair of toothed dentaries to the taxon, which Currie et al. (1990) made the holotype of Richardoestesia. Holtz (DML, 1998) noted the holotype is labeled "Steneodactylus pergracilis" in the CMN's collection, showing that was an earlier proposed name for the taxon. This has not been published in the literature however. Sternberg later (1932) described Macrophalangia canadensis based on a pes from the same formation found in 1928, then thought to be an ornithomimid. Colbert and Russell (1969) noted the two forms may be synonymous, though this was not shown to be likely until Osmólska (1981) described Elmisaurus which preserved a manus and pes of similar morphology. An alternative hypothesis, the synonymy of Chirostenotes with Dromaeosaurus, was suggested by Ostrom (1969; 1990), but is clearly incorrect given Dromaeosaurus' preserved metacarpal I and the recently described partial skeleton UALVP 59400. Macrophalangia is here provisionally referred to Caenagnathus collinsi, as in Longrich et al. (2013).
Currie and Russell (1988) first suggested Chirostenotes was synonymous with the oviraptorosaurian mandible Caenagnathus, which was strengthened by Sues' (1994) Horseshoe Canyon specimen with Chirostenotes-like postcrania and an edentulous maxilla and shown to be true at the family level by Anzu (Lamanna et al., 2011) which preserved skeletons with Caenagnathus-like mandibles. As noted above, Longrich et al. (2013) separate Caenagnathus collinsi from Chirostenotes pergracilis based on differences between the type mandible of the former and Caenagnathus sternbergi, which they synonymized with pergracilis (as partial skeleton UALVP 59400 later confirmed). Cracraft (1971) named Caenagnathus sternbergi, known from a posterior mandible that differs from C. collinsi in a few characters. Currie et al. (1993) later described five dentaries which also differ from C. collinsi. They referred to these as Caenagnathus cf. sternbergi, as none are directly comparable to the C. sternbergi holotype. Currie (2005) illustrated a new mandible (TMP 2001.012.0012) which confirms Currie et al. were correct to refer the dentaries to C. sternbergi. This was described in detail by Funston and Currie (2014a). Longrich et al. referred TMP 1990.056.0006 to Chirostenotes, but 1979.008.0622, 1991.144.0001 and 1992.036.0390 to their Leptorhynchos elegans. Funston et al. (2019) reported histological work on dentaries which showed small cf. Leptorhynchos elegans UALVP 55639 was older than larger Chirostenotes TMP 1985.043.0070 and 1992.036.1237. They further referred TMP 1992.040.0044 to Chirostenotes, but Funston (2020) later placed this as cf. elegans instead. As UALVP 55639 with a 22.1 mm symphyseal length is completely remodeled and thus quite old, I have tentatively reassigned all larger dentaries (including TMP 1991.144.0001, 1992.036.0390 and 1992.040.0044) to Chirostenotes and smaller TMP 1979.008.0622 to Caenagnathidae indet. pending histological examination of each specimen. This is supported by Funston et al. reporting that fibrous external bone texture is present in histologically juvenile TMP 1985.043.0070 and also 1992.036.0390 and 1992.040.0044. Notably, the sternbergi holotype matches pergracilis TMP 2001.012.0012 in size, whereas it would be much smaller if it was a synonym of elegans.
TMP 1979.020.0001 was discovered in 1979 and described by Currie and Russell in 1988 as Chirostenotes pergracilis. They distinguished between two kinds of metatarsus (robust CMN 8538; gracile ROM 781 and TMP 1979.020.0001), tentatively believing them to be sexual morphs. Ironically, Currie and Russell had mentioned TMP 1982.039.0004 as an American Elmisaurus specimen although they did not yet refer the elegans holotype to that genus. Currie (1989) expanded on the separation, referring some specimens to small and fused Elmisaurus elegans (ROM 781, 37163 and TMP 1982.039.0004) and others to large and unfused Chirostenotes pergracilis (CMN 8538 and 9570, and TMP 1979.020.0001). Currie and Russell had proposed Caenagnathus sternbergi belonged to the elegans morph, but this was eventually disproven by Funston and Currie (2021) who described UALVP 59400, a partial skeleton combining a sternbergi-type mandible with a histologically adult Chirostenotes-type tarsometatarsus. Currie and Russell had referred TMP 1979.020.0001 to their elegans morph based on the slender third manual digit and gracile pes. They note the minimum width of manual phalanx III-3 is 10% of its length, compared to 11% in the Chirostenotes holotype and 10% in the Elmisaurus holotype. Sues (1997) followed this referral, but believed the gracile morph to only be a separate species, which he called Chirostenotes elegans. Currie and Russell described several characters consistant with TMP 1979.020.0001 being pergracilis instead of elegans- unfused tarsometatarsus; metatarsals II and IV much shorter than III (87 and 90% respectively); proximal end of metatarsal III diamond-shaped. These characters are more numerous and observable in more specimens than the 1% difference in phalangeal width, while the gracile pes may be due to the small size of TMP 1979.020.0001 compared to CMN 8538. Sullivan et al. (2011) questioned the referral of TMP 1979.020.0001 to Chirostenotes pergracilis, based on "some minor morphological differences, including a broader curvature of unguals I and II, a deeper ungual (measured from the dorsal and plantar extremity of the articular facet as per Senter, 2007), and a shorter phalanx III-3." Yet such differences are common within theropod species known from multiple specimens. Thus TMP 1979.020.0001 is assigned to Chirostenotes pergracilis here, as it has been by other more recent authors. A potential complication was described by Simon and Evans (2021) who used histology to determine TMP 1979.020.0001 was a juvenile with two LAGS, despite being slightly larger than the histologically adult UALVP 59400. They suggested the former may be a juvenile Caenagnathus instead, but if this is true it could also be the case for the Chirostenotes holotype which has never been examined histologically, and could thus raise significant taxonomic issues. Currie and Russell referred manual ungual II TMP 1979.014.0499 to Chirostenotes, but it has not been described or figured. The size is identical to TMP 1979.020.0001 so it is tentatively retained in Chirostenotes here, although this would also make sense for a young Caenagnathus. Notably, differences between manual ungual II in Caenagnathus and Chirostenotes have since been discovered, so it should be possible to assign TMP 1979.014.0499 eventually.
Funston (2020) redescribed the holotype and TMP 1979.020.0001. Note that for the latter specimen, Funston lists manual ungual I as III in Table 2 and recognized distal tarsals Currie and Russel had not commented on. A phalanx III-3 and ungual III had been lost from the holotype, and manual phalanx III-1 from TMP 1979.020.0001. Funston described anterior dentaries TMP 1996.012.0142 as Caenagnathidae indet., but gievn their large size and distinction from Caenagnathus (short symphysis, upturned dentary tip), they are referred to Chirostenotes here. The partial skeleton UALVP 59400 was found on June 23 2016 and described as Chirostenotes pergracilis by Funston and Currie (2021). Note their Figure 3 has the articulated cervical vertebrae identified as 4, 5 and 6 but the text and Table 1 place them as 7,8 and 9. It is notable for combining mandibular and pedal material in the Dinosaur Park Formation, showing the sternbergi mandible type is associated with the tarsal morphology of Macrophalangia.
Varricchio (2001) described MOR 1107, an articular from the Two Medicine Formation, as Caenagnathus sternbergi. The specimen does have sternbergi's articular characters as diagnosed by Cracraft, so is here referred to Chirostenotes pergracilis which it is contemporaneous with. While Citipes does not preserve articular material, comparison with complete Chirostenotes mandible TMP 2001.012.0012 shows MOR 1107 is comparable in size and much larger than Citipes.
References- Gilmore, 1924. A new coelurid dinosaur from the Belly River Cretaceous Alberta. Canada Geological Survey, Bulletin 38, geological series 43, 1-13.
Sternberg, 1932. Two new theropod dinosaurs from the Belly River Formation of Alberta. Canadian Field-Naturalist. 46(5), 99-105.
Colbert and Russell, 1969. The small Cretaceous dinosaur Dromaeosaurus. American Museum Novitiates. 2380, 1-49.
Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Peabody Museum Bulletin. 30, 165 pp.
Cracraft, 1971. Caenagnathiformes: Cretaceous birds convergent in jaw mechanism to dicynodont reptiles. Journal of Paleontology. 45(5), 805-809.
Osmólska, 1981. Coossified tarsometatarsi in theropod dinosaurs and their bearing on the problem of bird origins. Palaeontologia Polonica. 42, 79-95.
Currie and Russell, 1988. Osteology and relationships of Chirostenotes pergracilis (Saurischia, Theropoda) from the Judith River (Oldman) Formation of Alberta, Canada. Canadian Journal of Earth Sciences. 25(7), 972-986.
Currie, 1989. The first records of Elmisaurus (Saurischia, Theropoda) from North America. Canadian Journal of Earth Sciences. 26(6), 1319-1324.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation of southern Alberta, Canada. In Carpenter and Currie (eds.). Dinosaur Systematics: Perspectives and Approaches. Cambridge University Press. 107-125.
Ostrom, 1990. Dromaeosauridae. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria. University of California Press. 269-279.
Currie, Godfrey and Nessov, 1993 (published 1994). New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30(10), 2255-2272.
Sues, 1994. New evidence concerning the phylogenetic position of Chirostenotes (Dinosauria: Theropoda). Journal of Vertebrate Paleontology. 14(3), 48A.
Sues, 1997. On Chirostenotes, a Late Cretaceous oviraptorosaur (Dinosauria: Theropoda) from western North America. Journal of Vertebrate Paleontology. 17(4), 698-716.
Holtz, DML 1998. https://web.archive.org/web/20210603185802/http://dml.cmnh.org/1998Aug/msg00743.html
Snively, Currie, Brinkman, Ryan, Braman, Gardner, Lam, Spivak and Neuman, 2001. Alberta's dinosaurs and other fossil vertebrates: Judith River and Edmonton groups (Campanian-Maastrichtian). In Hill (ed.). Mesozoic and Cenozoic Paleontology in the western plains and Rocky Mountains. Guidebook for the Field Trips of the Society of Vertebrate Paleontology 61st Annual Meeting. Museum of the Rockies Occasional Paper No. 3. 47-75.
Varricchio, 2001. Late Cretaceous oviraptorosaur (Theropoda) dinosaurs from Montana. In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life. Indiana University Press. 42-57.
Currie, 2005. Theropods, including birds. In Currie and Koppelhus (eds.). Dinosaur Provincial Park, a spectacular ecosystem revealed. Part Two, Flora and Fauna from the park. Indiana University Press. 367-397.
Senter and Parrish, 2005. Functional analysis of the hands of the theropod dinosaur Chirostenotes pergracilis: Evidence for an unusual paleoecologial role. PaleoBios. 25(2), 9-19.
Lamanna, Sues, Schachner and Lyson, 2011. A new caenagnathid oviraptorosaur (Theropoda: Maniraptora) from the Upper Cretaceous (Maastrichtian) Hell Creek Formation of the western United States. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 140.
Sullivan, Jasinski and van Tomme, 2011. A new caenagnathid Ojoraptorsaurus boerei, n. gen., n. sp. (Dinosauria, Oviraptorosauria), from the Upper Cretaceous Ojo Alamo Formation (Naashoibito Member), San Juan Basin, New Mexico. New Mexico Museum of Natural History and Science Bulletin. 53, 418-428.
Longrich, Barnes, Clark and Millar, 2013. Caenagnathidae from the Upper Campanian Aguja Formation of west Texas, and a revision of the Caenagnathinae. Bulletin of the Peabody Museum of Natural History. 54(1), 23-49.
Funston and Currie, 2014a. A previously undescribed caenagnathid mandible from the late Campanian of Alberta, and insights into the diet of Chirostenotes pergracilis (Dinosauria: Oviraptorosauria). Canadian Journal of Earth Sciences. 51(2), 156-165.
Funston and Currie, 2014b. New Elmisaurus (Dinosauria: Oviraptorosauria) material from Mongolia and Alberta, Canada, and its bearing on North American caenagnathid taxonomy. Journal of Vertebrate Paleontology, Program and Abstracts, 2014. 134.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica Polonica. 61(1), 159-173.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria (Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Funston, 2020. Caenagnathids of the Dinosaur Park Formation (Campanian) of Alberta, Canada: Anatomy, osteohistology, taxonomy, and evolution. Vertebrate Anatomy Morphology Palaeontology. 8, 105-153.
Funston, Wilkinson, Simon, Leblanc, Wosik and Currie, 2020 (online 2019). Histology of caenagnathid (Theropoda, Oviraptorosauria) dentaries and implications for development, ontogenetic edentulism, and taxonomy. The Anatomical Record. 303(4), 918-934.
Funston and Currie, 2021 (online 2020). New material of Chirostenotes pergracilis (Theropoda, Oviraptorosauria) from the Campanian Dinosaur Park Formation of Alberta, Canada. Historical Biology. 33(9), 1671-1685.
Simon and Evans, 2021. Osteohistology supports immature ontogenetic status of North American oviraptorosaurs Apatoraptor pennatus and Chirostenotes pergracilis. The Society of Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual Meeting. 236-237.

Elmisaurinae Osmólska, 1981 vide Currie, 2000
Definition- (Elmisaurus rarus <- Caenagnathus collinsi) (Hendrickx, Hartman and Mateus, 2015)
References- Osmólska, 1981. Coossified tarsometatarsi in theropod dinosaurs and their bearing on the problem of bird origins. Palaeontologia Polonica. 42, 79-95.
Currie, 2000. Theropods from the Cretaceous of Mongolia. The Age of Dinosaurs in Russia and Mongolia. 434-455.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.

unnamed elmisaurine (Funston, Currie and Burns, 2016)
Late Maastrichtian, Late Cretaceous
Frenchman Formation, Saskatchewan, Canada
Material- (RSM P2600.1) ?scapula, proximal tibia, distal tibia, proximal fibula, astragalocalcaneum, distal metatarsal II, distal metatarsal III (~15.8 mm trans)
Comments- RSM P2600.1 was found in 1989 and described as Leptorhynchos sp. (a genus in which they included what is now Citipes) by Funston et al. (2016). They noted "posterior surface has two cruciate ridges" as in Elmisaurus and Citipes (also now known in Anzu), but claimed it differed from elegans in that the latter has distal metatarsal III deeper than wide. However all Citipes elegans have a surface wider than deep as in RSM P2600.1. The strongly flaring distal tibia (176% of shaft width) is most similar to Elmisaurus, while the age matches Elmisaurus, Eoneophron and Anzu. Differences from Eoneophron noted by Atkins-Weltman et al. (2024) include- more developed postfibular flange on tibia; astragalocalcanear condyles subequal in size and distal extent; narrower transverse expansion of metatarsal III midshaft; distal end of metatarsal III deeper compared to width.
References- Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica Polonica. 61(1), 159-173.
Atkins-Weltman, Simon, Woodward, Funston and Snively, 2024. A new oviraptorosaur (Dinosauria: Theropoda) from the end-Maastrichtian Hell Creek Formation of North America. PLoS ONE. 19(1): e0294901.

unnamed elmisaurine (Varricchio, 2001)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Dawson County, Montana, US
Material- (MOR 752) astragalar fragment, partial metatarsal II (~131 mm), distal phalanx II-1, phalanx II-2 (26.1 mm), metatarsal fragment, phalanx III-1 (34 mm), phalanx III-2 (23.3 mm), phalanx III-3 (25.1 mm), pedal ungual III (24.3 mm), phalanx IV-1 (23.1 mm), phalanx IV-2 (16.2 mm), phalanx IV-3 (14.4 mm), phalanx IV-4 (16.5 mm), pedal ungual IV (21 mm)
Comments- Varricchio (2001) referred a pes (MOR 752) to Elmisaurus elegans (now Citipes). This was based on lacking the long phalanges of Chirostenotes, being more slender than Elmisaurus rarus, and possessing "a small but distinct anterolateral process just proximal to the distal articulation" of metatarsal II. The latter two characters have proven to not separate the taxa once more specimens of each were recovered (see Other diagnoses under Citipes). However the dorsoventral expansion of distal condyles on pedal phalanges and short phalanx III-2 are similar to Elmisaurus rarus (ZPAL MgD-I/98) instead of Citipes elegans (TMP 2000.012.0008), and indeed the shapes of phalanges II-1, III-1 and III-2 are nearly identical. Funston et al. (2016) provisionally retained it in elegans, although they noted that elegans specimen TMP 2000.012.0008 lacks a phalanx III-3 that is longer than III-2 as seen in MOR 752. This is also unlike Elmisaurus, but intriguely present in Macrophalangia (= Caenagnathus?). Atkins-Welman et al. (2024) suggested MOR 752 "may be more
likely" to be a distinct taxon from the larger Eoneophron and Anzu based on the anterior process on metatarsal II (inferred to be absent in Eoneophron based on the lack of a process on its metatarsal IV), which would be expected to be better developed in older individuals. As noted however this process is individually variable in Citipes, absent in the relatively large TMP 2000.012.0008, so MOR 752 and Eoneophron cannot be directly compared. Both are from the same formation and differ from Elmisaurus and Citipes, so could very well belong to the same taxon but more complete remains will be necessary to confirm this.
References- Varricchio, 2001. Late Cretaceous oviraptorosaur (Theropoda) dinosaurs from Montana. In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life. Indiana University Press. 42-57.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica Polonica. 61(1), 159-173.

Eoneophron Atkins-Weltman, Simon, Woodward, Funston and Snively, 2024
E. infernalis Atkins-Weltman, Simon, Woodward, Funston and Snively, 2024
Etymology- "Genus name derived from the Ancient Greek "eo"–meaning "dawn," and from the genus name of the Egyptian vulture, Neophron, sometimes referred to as the "pharaoh’s chicken." The species name derives from Latin for Hell, in reference to the Hell Creek Formation. Together the taxon name equates to "Pharaoh's dawn chicken from Hell.""
Late Masstrichtian Late Cretaceous
Hell Creek Formation, Meade County, South Dakota, US
Holotype- (CM 96523) (78 kg using Campione et al.; 6 year old subadult) incomplete right femur (390 mm), right tibiotarsus (499 mm), right metatarsal III (247 mm), right distal tarsal IV fused to metatarsal IV (233 mm)
Diagnosis- (after Atkins-Weltman et al., 2024; autapomorphies only) astragalocalcaneum fused to tibia; shaft of metatarsal IV with well-developed oblique longitudinal ridge on anterior surface extending along distal three-quarters of shaft.
Comments- The holotype was purchased in 2020 and initially identified as Anzu (Pester, 2024 online). Note Figure 1 incorrectly labels the femur the left element. Atkins-Weltman et al. (2024) find "the presence of a putative neonatal line followed by six LAGs in metatarsal IV suggests the death of this individual occurred during its sixth year of life. Because the sixth LAG is so close to the periosteal surface and often merges with the surface in each bone examined, death probably occurred shortly after this individual emerged from its annual growth hiatus." Furthermore, "the zones between the closely spaced outermost LAGs in metatarsal IV ... remain vascularized with high osteocyte lacuna density, suggesting that this larger metatarsal was approaching asymptotic size but was still growing in length at death."
Atkins-Weltman et al. (2024) recovered Eoneophron in a polytomy including all caenagnathids except Microvenator and Gigantoraptor (though only a majority rule consensus is shown), but excluding largely incomparable taxa (Apatoraptor, Caenagnathasia, Epichirostenotes) a posteriori reveals it is the sister taxon to Citipes plus Elmisaurus.
References- Atkins-Weltman, Simon, Woodward, Funston and Snively, 2024. A new oviraptorosaur (Dinosauria: Theropoda) from the end-Maastrichtian Hell Creek Formation of North America. PLoS ONE. 19(1): e0294901.
Pester, 2024 online. 'I felt my heart skip a beat': Researcher discovers dinosaur 'chicken from hell' after buying fossil online. Live Science. January 24.

Elmisaurus Osmólska, 1981
Other diagnoses- Osmólska (1981) listed proximolateral process on metatarsal IV as diagnostic at genus level, but it is also present in Chirostenotes, Eoneophron and Citipes and has not been shown to be absent in any caenagnathid.
Currie (1989) stated metatarsal II is subequal in length to III (>93%) in both Elmisaurus and Citipes (his E. elegans), but this is not true in some Citipes specimens (II is 89% in TMP 1982.016.0006 and TMP 2000.012.0008) and Avimimus shows this as well (93%). Similarly, IV being close to III in length is true of all caenagnathids where known (Eoneophron, Elmisaurus, Citipes, Kol, Avimimus) except Chirostenotes. Metatarsal II curving anterodistally is present in all caenagnathids. While Currie stated the proximal end of metatarsal III is triangular in Elmisaurus, unlike Chirostenotes, the latter does have a triangular section at some points. The only differences in cross sectional shape between caenagnathid taxa in the literature is due to proximodistal position along the metatarsal (see detailed discussion under Leptorhynchos comments).
Currie et al. (2016) listed the "large, compound proximal protuberance on the posterior surfaces of metatarsals II-IV" as being diagnostic for elmisaurines including Elmisaurus and Citipes, but this is present in Caenagnathus (based on mtII) and Chirostenotes as well. They also proposed better development of m. tibialis cranialis tubercles on the dorsal surface of metatarsals II-IV differed from Chirostenotes, but while these are present in the proximal middle section of II-IV Elmisaurus' and Citipes' holotypes, variation exists in other specimens. Elmisaurus IGM 102/6 has them on III and IV but not II, Elmisaurus IGM 102/7 lacks them on II and IV at least, and Citipes TMP 2000.012.0008 lacks them on III and IV at least. Confusingly, Currie et al. also refer to scars on the distal ends of metarsals as being these insertions (e.g. Figure 9A1), which is the only place Funston et al. (2016) describe them as being. Such distal scars must be very faint however, since they were not illustrated in either holotype and are not obvious in most photos either. Due to this, they are not used in diagnoses here.
References- Osmólska, 1981. Coossified tarsometatarsi in theropod dinosaurs and their bearing on the problem of bird origins. Palaeontologia Polonica. 42, 79-95.
Currie, 1989. The first records of Elmisaurus (Saurischia, Theropoda) from North America. Canadian Journal of Earth Sciences. 26(6), 1319-1324.
Currie, Funston and Osmólska, 2016 (online 2015). New specimens of the crested theropod dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica. 61(1), 143-157.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica Polonica. 61(1), 159-173.
E. rarus Osmólska, 1981
= Chirostenotes rarus (Osmólska, 1981) Paul, 1988
Early Maastrichtian, Late Cretaceous
Nemegt, Nemegt Formation, Mongolia
Holotype- (ZPAL MgD-I/172) (1.68 m) tarsometatarsus (163 mm; II 147, III 157, IV 147 mm)
Paratypes- (ZPAL MgD-I/20) proximal tarsometatarsus (mtIII ~173 mm)
(ZPAL MgD-I/98) rib fragments, gastralial fragments, two partial mid sacral centra (37 mm), proximal scapula, metacarpal I (45 mm), phalanx I-1 (65 mm), manual ungual I (44 mm), incomplete metacarpal II (~63 mm), phalanx II-1 (66 mm), phalanx II-2 (66 mm), phalanx III-1 (30 mm), phalanx III-2 (30 mm), phalanx III-3 (43 mm), proximal manual ungual, proximal pubes, partial ischia, partial femora (~308 mm), partial tibia (~340 mm), phalanx I-1 (26 mm), pedal ungual I (~23 mm), distal metatarsal II, phalanx II-1 (45 mm), partial phalanx II-2 (34 mm), fragmentary pedal ungual II (~31 mm), distal metatarsal III (~151 mm), phalanx III-1 (46 mm), phalanx III-2 (32 mm), phalanx III-3 (30 mm), distal metatarsal IV, fragmentary phalanx IV-1, phalanx IV-2 (22 mm), phalanx IV-3 (19 mm), phalanx IV-4 (18 mm), fragmentary pedal ungual IV (~26 mm), fragments
Referred- (IGM 102/6; field number PJC 2000.1) tarsometatarsus (194 mm; II 172.4, III 185, IV 175.7 mm), metatarsal V (70.3 mm) (Currie, 2001; described by Currie, Funston and Osmólska, 2016)
(IGM 102/7; field number PJC 2000.2) (~23 kg adult) frontal (50.5 mm), ~third to fifth cervical neural arch, partial posterior cervical vertebra (26.4 mm), incomplete anterior dorsal vertebra (27.7 mm), three proximal dorsal ribs, dorsal rib fragments, gastralial fragments, vertebral fragment, distal phalanx II-2, incomplete manual ungual II (40 mm), phalanx III-1 (28 mm), phalanx III-3 (40 mm), proximal femur (~246 mm), tibiae (320, 327 mm), pedal phalanx I-1 (23.1 mm), incomplete pedal ungual I (30 mm), distal tarsal III, (tarsometatarsus ~176 mm) metatarsal II (161.8 mm), partial metatarsal III (~172 mm), metatarsal IV (162.2 mm) (Currie, 2001; described by Currie, Funston and Osmólska, 2016)
(IGM 102/8; field number PJC 2000.3) metatarsal IV (164 mm) (Currie, 2001; described by Currie, Funston and Osmólska, 2016)
(IGM 102/9; field number PJC 2001.8) proximal tarsometatarsus (Currie, Funston and Osmólska, 2016)
(IGM 102/10; field number PJC 2002.4) vertebra, tibia (Currie, Funston and Osmólska, 2016)
?(?IGM coll.) femur (Currie, 2001)
Early Maastrichtian, Late Cretaceous
Bugin Tsav, Nemegt Formation, Mongolia
?(IGM 102/107) anterior dentaries (symph 20.63 mm) (Tsuihiji, Watabe, Tsogtbaatar and Barsbold, 2016)
?(IGM coll.; field number NatGeo.2018.040) partial ?postorbital, dentary fragment, partial angular, ?anterior cervical vertebral fragment, partial scapula, coracoid, pubes (one incomplete, one fragmentary), partial astragalus, proximal metatarsal III, incomplete metatarsal IV, metatarsal fragments (Funston, 2019)
? Early Maastrichtian, Late Cretaceous
? Nemegt Formation, Mongolia
? ventral skull (www.paleofile.com)
Diagnosis- (after Osmólska, 1981) metacarpal I ~66% [69%] the length of phalanx I-1 (76% in Hagryphus; 50% in Apatoraptor; but unknown in other caenagnathids); tarsometatarsal fusion (also in Citipes; unknown in other caenagnathids except Anzu, Caenagnathus, Chirostenotes and Eoneophron); posterior surface of metatarsus deeply concave (also in Anzu and Citipes; unknown in other caenagnathids except Caenagnathus and Chirostenotes); length/width ratio of metatarsus 0.18 (also in Citipes; unknown in other caenagnathids except Anzu, Caenagnathus? and Chirostenotes).
(after Currie et al., 2016) posterior medial and lateral ridges of metatarsal III midshaft close to trochlear ridges (also in Citipes and Eoneophron; unknown in other caenagnathids except Anzu and Chirostenotes).
Other diagnoses- Osmólska (1981) listed several supposedly diagnostic characters- digit III markedly thinner than digits I and II both of which are equally thick (untrue as digit I is more gracile than II, and conditions are the same in other caenagnathids); ventroposterior portions of manual phalanges distinctly thickened at proximal surfaces (also in other caenagnathids); metatarsal III visible anteriorly for ~90% of its length (also in the Citipes holotype, but not in Elmisaurus IGM 102/7).
Currie et al. (2016) stated at least manual unguals I and II of Elmisaurus (III is unknown) were more robust and curved than Chirostenotes, with lower proximodorsal lips, but they are comparable to ungual I of Caenagnathus, Hagryphus and Apatoraptor (II unknown except for Apatoraptor).
(suggested) ventral edge of ischium convex distal to obturator process.
Comments- Discovered in 1970, listed as "small theropod gen. et sp. nov." by Osmólska (1980) and described in 1981, Elmisaurus rarus was the first caenagnathid discovered with both manual and pedal remains. This allowed the synonymy between Chirostenotes and Macrophalangia to be demonstrated. Elmisaurus has been viewed as a relative of Chirostenotes (first in Elmisauridae and later in Caenagnathidae) from its discovery until the 2000s, when this was questioned by Maryanska et al. (2002). They noted it differs from Chirostenotes in having a vascular foramen between metatarsals III and IV, an m. tibialis cranialis tubercle on the dorsal surfaces of metatarsals II-IV, a deeply concave posterior side, and a proximolateral process on metatarsal IV. Yet these are all apomorphies that tell us nothing about Elmisaurus' relationships unless we find non-caenagnathid taxa that share them. Maryanska et al. stated pygostylians have the first two characters, and Avimimus the last, but Elmisaurus otherwise resembles oviraptorosaurs and is quite dissimilar to any paravian. The relationship with Avimimus is possible, as found in my modified version of the Hartman et al. matrix, but that taxon is a caenagnathid in that tree anyway. It is especially confusing that Maryanska et al. continue to refer Citipes to Caenagnathidae, as it shares the characters they describe for Elmisaurus. Citipes is similar enough to Chirostenotes pergracilis to be synonymized by some authors (e.g. Currie and Russell, 1988), and Elmisaurus has been synonymized with Chirostenotes by others (e.g. Paul, 1988). It is completely unwarranted to widely separate the two genera, and Elmisaurus has more recently been reestablished as a caenagnathid (e.g. Longrich et al., 2013; Currie et al., 2016).
Currie (2001, 2002) first mentioned new specimens of Elmisaurus rarus, most of which were later noted in an abstract (Funston and Currie, 2014). Currie et al. (2016) described these specimens, as well as additional material of ZPAL MgD-I/98. They identified the manual ungual of the latter as "almost certainly" from digit I, which was the possible identification of Osmólska as well despite her figure 2 placing it on digit II. While Funston and Currie state the frontal "suggests the presence of a high crest", which is also assumed by Currie et al., no evidence is presented to support this and the frontal shows no dorsal projection. Only one specimen mentioned by Currie (2001/2002) was not described in Currie et al.- a femur referred to cf. Elmisaurus rarus. Sniovely (2000) uses a metatarsus of "Elmisaurus sp. (TMP:PJC}" in his thesis, which based on the outline in his Figure 2.11 is the holotype or a cast of it. Tsuihiji et al. (2016) described a partial mandible that is here referred to E. rarus based on stratigraphy, but it cannot be distinguished from Chirostenotes based on morphology. Ford (2015 online) reported a partial skull photographed in "a Japanese guide book".
References- Osmólska, 1980. The Late Cretaceous vertebrate assemblages of the Gobi Desert, Mongolia. Memoires de la Societe Geologique de France. 139, 145-150.
Osmólska, 1981. Coossified tarsometatarsi in theropod dinosaurs and their bearing on the problem of bird origins. Palaeontologia Polonica. 42, 79-95.
Currie and Russell, 1988. Osteology and relationships of Chirostenotes pergracilis (Saurischia, Theropoda) from the Judith River (Oldman) Formation of Alberta, Canada. Canadian Journal of Earth Sciences. 25(7), 972-986.
Paul, 1988. The Predatory Dinosaurs of the World. Simon and Schuster. 464 pp.
Snively, 2000. Functional morphology of the tyrannosaund arctometatarsus. Masters Thesis, University of Calgary. 273 pp.
Currie, 2001. Nomadic Expeditions, Inc. report on fieldwork in Mongolia, September 2000. Alberta Palaeontological Society, Fifth Annual Symposium. 12-16.
Currie, 2002. Report on fieldwork in Mongolia, September 2001. Alberta Palaeontological Society, Sixth Annual Symposium. 8-12.
Maryanska, Osmólska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Longrich, Barnes, Clark and Millar, 2013. Caenagnathidae from the Upper Campanian Aguja Formation of West Texas, and a revision of the Caenagnathinae. Bulletin of the Peabody Museum of Natural History. 54(1), 23-49.
Funston and Currie, 2014. New Elmisaurus (Dinosauria: Oviraptorosauria) material from Mongolia and Alberta, Canada, and its bearing on North American caenagnathid taxonomy. Journal of Vertebrate Paleontology, Program and Abstracts, 2014. 134.
Ford, 2015 online. http://www.paleofile.com/Dinosaurs/Theropods/Elmisaurussp.asp
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of Alberta, Canada. Cretaceous Research. 54, 179-187.
Currie, Funston and Osmólska, 2016 (online 2015). New specimens of the crested theropod dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica. 61(1), 143-157.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica Polonica. 61(1), 159-173.
Tsuihiji, Watabe, Tsogtbaatar and Barsbold, 2016. Dentaries of a caenagnathid (Dinosauria: Theropoda) from the Nemegt Formation of the Gobi Desert in Mongolia. Cretaceous Research. 63, 148-153.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria (Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.

Citipes Funston, 2020
C. elegans (Parks, 1933) Funston, 2020
= Ornithomimus elegans Parks, 1933
= Macrophalangia elegans (Parks, 1933) Koster, Currie, Eberth, Brinkman, Johnston and Braman, 1987
= Chirostenotes elegans (Parks, 1933) Currie and Russell, 1988
= Elmisaurus elegans (Parks, 1933) Currie, 1989
= "Leptorhynchos" elegans (Parks, 1933) Longrich, Barnes, Clark and Millar, 2013a
= Leptorhynchos elegans (Parks, 1933) Longrich, Barnes, Clark and Millar, 2013b
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada
Holotype- (ROM 781) partial distal tarsal III, distal tarsal IV, metatarsal II (155 mm), partial metatarsal III (161 mm), metatarsal IV (157 mm)
Referred- (ROM 37163) distal metatarsal II (Currie, 1989)
?(TMP 1981.023.0034) partial ilium (Rhodes, Funston and Currie, 2020)
....(TMP 1981.023.0035) partial ilium (Rhodes, Funston and Currie, 2020)
....(TMP 1981.023.0039) last sacral vertebra (Funston, 2020)
(TMP 1982.016.0006) tarsometatarsus (mtII 152.4, mtIII 172.2, mtIV 160.5, mtV 44.3 mm) (Funston et al., 2016)
(TMP 1982.039.0004) proximal tarsometatarsus (Currie, 1989)
?(TMP 1984.163.0036) distal metatarsal III (Funston et al., 2016)
?(TMP 1986.036.0186) distal metatarsal III (Funston et al., 2016)
(TMP 1988.036.0104) distal metatarsal II (Funston et al., 2016)
?(TMP 1992.036.0674) partial ilium (Funston, 2020)
?(TMP 1993.036.0630) distal metatarsal III (Funston et al., 2016)
?(TMP 1994.012.0880) tibia (279.66 mm) (Funston et al., 2016)
(TMP 1996.012.0141) incomplete tarsometatarsus (mtII ~130 mm, mtIV ~135 mm) (Currie, 2005; described by Funston et al., 2016)
(TMP 2000.012.0008) (tarsometatarsus ~185 mm) distal tarsal III fused to metatarsal II (164.8 mm), phalanx II-1, pedal ungual II, distal tarsal IV fused to metatarsal III (185.3 mm), phalanx III-1 (~44 mm), phalanx III-2, phalanx III-3, pedal ungual III, phalanx IV-?2 (22.3 mm) (Funston et al., 2016)
?(TMP 2005.049.0190) metatarsal III (Funston et al., 2016)
(UALVP 55585) (subadult) partial metatarsal III (Funston et al., 2016)
(UALVP 55639) (adult) anterior dentaries (symph 22.1 mm) (Funston, Wilkinson, Simon, Leblanc, Wosik and Currie, 2020)
(UALVP 59606) (6 year old adult) distal tarsal IV, metatarsal IV (146 mm) (Funston, 2020)
Late Campanian, Late Cretaceous
Aguja Formation, Texas, US
 ?(TMM 43057-36; paratype of Leptorhynchos gaddisi) distal tarsal IV fused to proximal metatarsal IV (~22.6 mm trans) (Longrich, Barnes, Clark and Millar, 2013a)
Diagnosis- (after Funston, 2020) distal tarsals III and IV coossified with each other and proximal metatarsus at maturity (also in Elmisaurus; unknown in other caenagnathids except Anzu, Caenagnathus and Chirostenotes); metatarsal III with prominent cruciate ridges on posterior surface (also in Anzu, Eoneophron and Elmisaurus; unknown in other caenagnathids except Chirostenotes).
(proposed) distal tibia flared transversely <150% of shaft width (also in Eoneophron; unknown in other caenagnathids except Anzu, Chirostenotes and Elmisaurus); distal condyles of pedal phalanges (II-1, III-1 and III-2 at least) less dorsoventrally expanded than Elmisaurus (unknown in other caenagnathids except Chirostenotes); pedal phalanx III-2 elongate (posterior height <50% of length between condyles and cotyles) (unknown in other caenagnathids except Caenagnathus? and Elmisaurus).
Other diagnoses- Currie (1989) proposed "close to the distal articular surfaces, small processes of metatarsals II and IV overlap metatarsal III" is diagnostic of elegans, but they are not present in TMP 2000.012.0008. Currie used the character "posteromedial corner of the tarsometatarsus is more deeply emarginated than that of Elmisaurus rarus" to distinguish elegans based on the rarus holotype versus TMP 1982.039.0004, but the deep emargination is also seen in Elmisaurus rarus specimen IGM 102/6 and not in elegans specimens ROM 781, TMP 1982.016.0006 or TMP 1996.012.0141. Currie stated metatarsal II was more slender in Citipes (maximum shaft diameter 9% of metatarsal length in holotype) than Elmisaurus (12% in holotype), but Elmisaurus rarus specimen IGM 102/6 has a ratio of 8%. He also listed "longitudinal, ridge-like posterolateral margin of metatarsal IV is not as powerfully developed proximally as that of Elmisaurus rarus", but while this appears true in his drawing of the elegans type, the ridge seems as well developed in all preserved areas although far proximally it is restored in plaster (Funston et al., 2016: Fig. 3C3).
Longrich et al. (2013a) proposed several differences in beak shape from Chirostenotes based on the assumption TMP 1992.036.0390 belonged to elegans- tip of beak strongly upturned, with anterior occlusal margin projecting vertically; anterior margin of symphysis straight; chin squarish in lateral view. Here 1992.036.0390 is referred to Chirostenotes (see below), so that these are individual variation and/or taphonomic.
Funston et al. (2016) claimed that "on metatarsal III of Elmisaurus rarus, there is rugosity on the distal base of the lateral cruciate ridge, and a small longitudinal ridge that bisects the longitudinal sulcus between the cruciate ridges proximally" and that "both of these features are absent in Leptorhynchos elegans." Yet these are not visible in Elmisaurus specimens IGM 102/6 or 102/7 (Fig. 10 of Currie et al., 2016) or available images of the holotype, so are not used here. Funston et al. also proposed the posterior extension of the proximal tarsometatarsus is smaller than Elmisaurus, but this is not true in TMP 2000.012.0008. They also claim in elegans "the distal third of the shaft of metatarsal II curves medially", but this is only true in TMP 2000.012.0008 and not the holotype. Similarly they say "metatarsal IV is straight along its length", but this is untrue in the later described UALVP 59606. Finally, they claim the "distal condyles of metatarsal III are deeper anteroposteriorly than wide mediolaterally" in elegans, but show the opposite in their Table 1 for TMP 1982.016.0006 and 2000.012.0008, which is the case in the holotype as well (Currie, 1989: Fig. 2r).
Comments- The holotype was discovered in 1926 and originally described as a species of Ornithomimus (Parks, 1933), though Sternberg (1934) soon recognized it was not an ornithomimid. Russell (1972) synonymized it with Macrophalangia canadensis, while Currie and Russell (1988) synonymized both with Chirostenotes pergracilis. The latter authors believed the elegans specimen to be a gracile morph of the species, which could be called Chirostenotes elegans if it was in fact taxonomically distinct. Currie (1989) described two new specimens (ROM 37163 and TMP 1982.039.0004), noting similarities to Elmisaurus rarus that were not seen in Chirostenotes pergracilis. He made the new combination Elmisaurus elegans. These similarities were said to be insufficient by Sues (1997) (without justification), who called the species Chirostenotes elegans. Maryanska et al. (2002) and Osmólska et al. (2004) also assigned elegans to Chirostenotes instead of Elmisaurus, though they never state their rationale.
When it came to cranial material, Currie and Russell (1988) first proposed the synonymy of Caenagnathus collinsi with Chirostenotes pergracilis, and Caenagnathus sternbergi with Citipes (then Chirostenotes) elegans. The new partial skeleton UALVP 59400 instead shows sternbergi mandibles went with pergracilis tarsometatarsi. Longrich et al. (2013a) placed elegans in their new genus Leptorhynchos, based on the Aguja Formation dentaries L. gaddisi. While the gaddisi and elegans material is similar in size and tarsometatarsal fusion, their reasons for separating these from Elmisaurus are flawed (see Leptorhynchos comments), and there was no reason to refer elegans to Leptorhynchos over Elmisaurus or vice versa. Funston and Currie (2014b) agreed and called it Elmisaurus elegans, but Currie et al. (2016) and Funston et al. (2020) referred elegans to Leptorhynchos without justification. Funston (2020) solved the issue by giving elegans its own genus Citipes, which functions regardless of phylogeny. Notably, recently recovered specimens show proposed differences between tarsometatarsi of Citipes and Elmisaurus vary individually within these taxa (see Other diagnoses above). However, the tibiae and pedal phalanges are different showing along with stratigraphy that they are not synonymous.
Longrich et al. (2013a) referred dentaries TMP 1979.008.0622, 1991.144.0001 and 1992.036.0390 to their Leptorhynchos elegans, and Funston et al. (2020) referred the third specimen as well. This was based on putative differences in beak shape (see Other diagnoses above) of 1992.036.0390, but it is similar in size to RTMP 1990.056.0006 which these authors considered to be Chirostenotes, while Citipes metatarsi are much smaller than those of Chirostenotes. Furthermore, the large TMP 2001.012.0012 has an intermediate beak shape, and 1990.056.0006 is taphonomically deformed to be assymetrical which may have straightened its beak tip. Adding further evidence, Funston et al. (2020) examined dentaries histologically and found small (symphyseal length 22.1 mm) UALVP 55639 to be a mature adult. Thus Funston (2020) concluded the TMP dentaries "cannot be confidently referred to Citipes elegans without osteohistological analysis showing that they are adults" and assigned them all to Caenagnathidae indet.. I've provisionally referred them all to Chirostenotes here, as all but one are larger than the confirmed mature UALVP 55639, two have fibrous external bone texture and two have poorly developed m. genioglossus attachments, both characters Funston et al. observed in histologically juvenile Chirostenotes TMP 1985.043.0070. This leaves UALVP 55639 the only recognized Citipes cranial element which is unfortunate as it is quite fragmentary. Funston and Currie (2014b) announced several new specimens in an SVP abstract, later describing them in Funston et al. (2016). One of these, partial tarsometatarsus TMP 1993.036.0181, was later reidentified by Funston and Currie (2021) as Chirostenotes pergracilis based on its large size and lack of fusion. Funston (2020) refers a few new specimens to Citipes- ilia and associated sacral TMP 1981.023.0034, 0035 and 0039, ilium TMP 1992.036.0674 and metatarsal IV UALVP 59606, describing them and figuring the metatarsal. The ilia and sacral are figured in Funston's (2019) thesis.
Note TMP 1992.036.0674 is incorrectly listed on the TMP online catalogue as Dromaeosauridae.
Here, the Leptorhynchos gaddisi holotype is considered indeterminate and possibly juvenile (see entry). However, among the referred material from the same formation (Longrich et al., 2013a) the distal tarsal IV and metatarsal IV (TMM 43057-36) are like Citipes and Elmisaurus in fusion and size. It's provisionally referred to the contemporeneous Citipes elegans here.
References- Parks, 1933. New species of dinosaurs and turtles from the Upper Cretaceous formations of Alberta. University of Toronto Studies, Geological Series. 34, 1-33.
Sternberg, 1934. Notes on certain recently described dinosaurs. Canadian Field Naturalist. 48, 7-8.
Cracraft, 1971. Caenagnathiformes: Cretaceous birds convergent in jaw mechanism to dicynodont reptiles. Journal of Paleontology. 45(5), 805-809.
Russell, 1972. Ostrich dinosaurs from the Late Cretaceous of western Canada. Canadian Journal of Earth Sciences. 9, 375-402.
Koster, Currie, Eberth, Brinkman, Johnston and Braman, 1987. Sedimentology and Palaeontology of the Upper Cretaceous Judith River/Bearpaw Formations at Dinosaur Provincial Park, Alberta, Field Trip #10. Geological Association of Canada, Mineralogical Association of Canada, Joint Annual Meeting, Saskatoon, Saskatchewan. 130 p.
Currie and Russell, 1988. Osteology and relationships of Chirostenotes pergracilis (Saurischia, Theropoda) from the Judith River (Oldman) Formation of Alberta, Canada. Canadian Journal of Earth Sciences. 25(7), 972-986.
Currie, 1989. The first records of Elmisaurus (Saurischia, Theropoda) from North America. Canadian Journal of Earth Sciences. 26(6), 1319-1324.
Currie, Godfrey and Nessov, 1993 (published 1994). New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30(10), 2255-2272.
Currie, 2005. Theropods, including birds. In Currie and Koppelhus (eds). Dinosaur Provincial Park, a spectacular ecosystem revealed, part two, flora and fauna from the park. Indiana University Press. 367-397.
Funston, Currie and Murray, 2013. Examining the diet of a toothless dinosaur: Evidence supporting a herbivorous diet in Caenagnathus (Dinosauria: Oviraptorosauria). Journal of Vertebrate Paleontology. Program and Abstracts 2013, 131.
Longrich, Barnes, Clark and Millar, 2013a. Caenagnathidae from the Upper Campanian Aguja Formation of west Texas, and a revision of the Caenagnathinae. Bulletin of the Peabody Museum of Natural History. 54(1), 23-49.
Longrich, Barnes, Clark and Millar, 2013b. Correction to "Caenagnathidae from the Upper Campanian Aguja Formation of west Texas, and a revision of the Caenagnathinae". Bulletin of the Peabody Museum of Natural History. 54(2), 263-264.
Funston and Currie, 2014a. A previously undescribed caenagnathid mandible from the late Campanian of Alberta, and insights into the diet of Chirostenotes pergracilis (Dinosauria: Oviraptorosauria). Canadian Journal of Earth Sciences. 51(2), 156-165.
Funston and Currie, 2014b. New Elmisaurus (Dinosauria: Oviraptorosauria) material from Mongolia and Alberta, Canada, and its bearing on North American caenagnathid taxonomy. Journal of Vertebrate Paleontology, Program and Abstracts, 2014. 134.
Currie, Funston and Osmólska, 2016 (online 2015). New specimens of the crested theropod dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica. 61(1), 143-157.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica Polonica. 61(1), 159-173.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria (Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Funston, 2020. Caenagnathids of the Dinosaur Park Formation (Campanian) of Alberta, Canada: Anatomy, osteohistology, taxonomy, and evolution. Vertebrate Anatomy Morphology Palaeontology. 8, 105-153.
Funston, Wilkinson, Simon, Leblanc, Wosik and Currie, 2020 (online 2019). Histology of caenagnathid (Theropoda, Oviraptorosauria) dentaries and implications for development, ontogenetic edentulism, and taxonomy. The Anatomical Record. 303(4), 918-934.
Rhodes, Funston and Currie, 2020. New material reveals the pelvic morphology of Caenagnathidae (Theropoda, Oviraptorosauria). Cretaceous Research. 114, 104521.

Avimimidae Kurzanov, 1981
Definition- (Avimimus portentosus <- Oviraptor philoceratops, Elmisaurus rarus, Caenagnathus collinsi) (Martyniuk, 2012)
Diagnosis- hyperarctometatarsus (unknown in Shixinggia).
References- Kurzanov, 1981. On the unusual theropods from Upper Cretaceous of Mongolia. Fossil Reptiles of Mongolia. Sovmestnaya Sovetsko-Mongol'skaya Paleontologicheskaya Ekspeditsiya, Trudy. 24, 39-50.
Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs. Vernon, New Jersey. Pan Aves. 189 pp.

Shixinggia Lu and Zhang, 2005
= "Shixinggia" Lu, 2004
S. oblita Lu and Zhang, 2005
= "Shixinggia oblita" Lu, 2004
Early Maastrichtian, Late Cretaceous
Zhenshui Formation, Nanxiong Group, Luyuan, Shixing County, Guangdong, China
Holotype- (BPV-112) eighth dorsal vertebra (25 mm), ninth dorsal vertebra (25 mm), tenth dorsal vertebra (25 mm), two incomplete dorsal ribs, sacrum (27, 27, 27, 30, 30, 30, 30 mm), first caudal vertebra, second caudal vertebra, third caudal vertebra, ilia (242 mm), proximal pubis, partial femur, proximal tibia, proximal fibula, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, phalanx IV-3, phalanx IV-4, pedal ungual IV
....(Shixing Museum coll.) fragments
Diagnosis- (modified from Lu and Zhang, 2005) preacetabular process lacking anteroventral process; anterioposteriorly shortened preacetabular process; large (pneumatic) foramen in the anterolateral surface of the proximal femur; small (pneumatic?) foramen in the proximomedial tibial surface.
Comments- This was collected in 1995 and mentioned as an oviraptorid in Lu et al.'s (2003) abstract. It was first named and described in Lu's (2004) thesis, then officially by Lu and Zhang (2005). While assigned to the Pingling Formation in botgh of those works, that has since been replaced by Li et al. 2007 by the Zhenshui Formation, Early Maastrichtian according to Xi et al 2021.
Lu and Zhang assign it to Oviraptoridae, though Lu (2004) found it to be a caenagnathid in a modified version of Maryanska's oviraptorosaur matrix, and sister taxon to Heyuannia within Oviraptoridae in a modified version of the TWiG matrix. Funston et al. (2016) and Hartman et al. (2019) recover it as sister to the Khaan+Heyuannia clade. Adding taxa to the latter analysis moves Shixinggia to Avimimidae.
References- Lu, Zhang and Li, 2003. A new oviraptorid dinosaur from the Late Cretaceous of Shixing, Nanxiong Basin of Guangdong Province, Southern China. Journal of Vertebrate Paleontology. 23(3), 73A.
Lu, 2004. Oviraptorid dinosaurs from southern China. PhD thesis, Southern Methodist University. 249 pp.
Lu, 2005. Oviraptorid dinosaurs from southern China. Geological Publishing House, Beijing. ISBN 7-116-04368-3. 200 pages + 8 plates.
Lu and Zhang, 2005. A new oviraptorid (Theropoda: Oviraptorosauria) from the Upper Cretaceous of the Nanxiong Basin, Guangdong Province of southern China. Acta Palaeontologica Sinica. 44(3), 412-422.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

Kol Turner, Nesbitt and Norell, 2009
K. ghuva Turner, Nesbitt and Norell, 2009
Late Campanian, Late Cretaceous
Ukhaa Tolgod, Djadokhta Formation, Mongolia
Holotype- (IGM 100/2011) distal tarsal, metatarsal I (17 mm), phalanx I-1 (21.7 mm), pedal ungual I, (metatarsus 225 mm) metatarsal II (203.5 mm), phalanx II-1 (44 mm), phalanx II-2 (29.9 mm), pedal ungual II, metatarsal III (109 mm), phalanx III-1 (43.2 mm), phalanx III-2 (31.2 mm), phalanx III-3 (24 mm), pedal ungual III (24.1 mm on curve), metatarsal IV (208.1 mm), phalanx IV-1 (23 mm), phalanx IV-2 (20.3 mm), phalanx IV-3 (16 mm), phalanx IV-4 (11.2 mm), pedal ungual IV (20 mm on curve), partial metatarsal V (~63 mm)
Diagnosis- (after Turner et al., 2009) combination of hyperarctometatarsus (also in Avimimus and derived alvarezsaurids) and robust flexor tubercles on pedal unguals (also in Patagonykus and other oviraptorosaurs).
Other diagnoses- Contra Turner et al.'s (2009) statement, metatarsal III does not extend over halfway up the metatarsus in Kol, extending 44% instead as in Avimimus. This is not higher up than other derived alvarezsaurids, as the ratio in Shuvuuia varies between 33% (IGM 100/1276) and 47% (IGM 100/975), though it is more than in Parvicursor and Ceratonykus. The short metatarsal II (98% of metatarsal IV length) is found in Avimimus, Parvicursor remotus and Ceratonykus as well, but not in Mononykus, Shuvuuia or IGM 100/99. Contary to Turner et al., Kol lacks deep extensor grooves on digit IV phalanges (pers. obs. IGM 100/2011). Contra their diagnosis, the accessory dorsomedial flange on metatarsal II they list is seemingly the dorsolateral flange on metatarsal IV, which is mentioned in the text as being absent in contrast to Mononykus and Shuvuuia. This is also present in Parvicursor and IGM 100/99, but absent in Alvarezsaurus and Avimimus, so is a symplesiomorphy.
Comments- Turner et al. (2009) described Kol as an alvarezsauroid, proposing it was more derived than Alvarezsaurus and Patagonykus, but outside the Shuvuuia plus Mononykus clade. It was later added to the TWiG analysis by Nesbitt et al. (2011) where it emerged in precisely this position. Agnolin et al. (2012) suggested Kol is actually a caenagnathid (related to Avimimus due to its hyperarctometatarsus), based on the prominently developed pedal flexor tubercles (plesiomorphic, also in Patagonykus), metatarsal II shorter than metatarsals III and IV (plesiomorphic, also in Alvarezsaurus, Linhenykus, Parvicursor and Ceratonykus; variable in Albinykus), lack of a dorsomedial flange on the medial side (plesiomorphic, also in Alvarezsaurus), and metatarsal III that extends more proximally than one-half of the total metatarsal length (plesiomorphic, also in Alvarezsaurus, Achillesaurus and Patagonykus). As can be seen, these characters might make Kol a comparatively basal alvarezsaurid, but they do not support an oviraptorosaurian identity any better. Hartman et al. (2019) found it more parsimonious for Kol to be a caenagnathid sister to Avimimus including all the character evidence of Turner et al., but an alvarezsauroid placement is only one step longer, suggesting more pedal characters must be analyzed.to support either position
References- Turner, Nesbitt and Norell, 2009. A large alvarezsaurid from the Late Cretaceous of Mongolia. American Museum Novitates. 3648, 14 pp.
Nesbitt, Clarke, Turner and Norell, 2011. A small alvarezsaurid from the eastern Gobi Desert offers insight into evolutionary patterns in the Alvarezsauroidea. Journal of Vertebrate Paleontology. 31(1), 144-153.
Agnolin, Powell, Novas and Kundrat, 2012 (online 2011). New alvarezsaurid (Dinosauria, Theropoda) from uppermost Cretaceous of north-western Patagonia with associated eggs. Cretaceous Research. 35, 33-56.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

Avimimus Kurzanov, 1981
Diagnosis- (after Kurzanov, 1981) frontal and parietal bones fused (unknown in other avimimids; also in Banji); last two cervical vertebrae with large hypapophyses (unknown in other avimimids); fibula highly reduced (unknown in other avimimids); fibula fused distally to tibiotarsus (unknown in Kol); metatarsals II and IV proximally fused (unknown in Shixinggia); fifth metatarsal fuses fully with the fourth (unknown in Shixinggia).
(after Funston et al., 2018) premaxilla invaded by antorbital fenestra (unknown in other avimimids); fused neurocranium (unknown in other avimimids); low mandible with poorly developed symphyseal shelf (unknown in other avimimids); fused tibiotarsus (unknown in other avimimids).
Other diagnoses- Kurzanov (1981) also included characters plesiomorphic for oviraptorosaurs- skull small with large orbits; additional femoral condyles for articulation with fibula; and common in other caenagnathoids- no fewer than 11 cervical vertebrae; cervical vertebrae increase in size posteriorly (e.g. in Apatoraptor); first three dorsal vertebrae with large hypapophyses; distal tarsals fused to metatarsals.
Funston et al. (2018) listed dorsal vertebrae without pleurocoels as diagnostic, but some dorsals have pleurocoels (Vickers-Rich et al., 2002). They also listed the hyperarctometatarsus, which is also present in Kol, and the lack of pedal digit I which is only known for A. portentosus.
References- Kurzanov, 1981. On the unusual theropods from Upper Cretaceous of Mongolia. In Resetov (ed.). Iskopaemye pozvonocnye Mongolii. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 15, 39-50.
Funston, Mendonca, Currie and Barsbold, 2018 (online 2017). Oviraptorosaur anatomy, diversity and ecology in the Nemegt Basin. Palaeogeography, Palaeoclimatology, Palaeoecology. 494, 101-120.
A. portentosus Kurzanov, 1981
Early Maastrichtian, Late Cretaceous
Yagaan Khovil, Nemegt Formation, Mongolia
Holotype- (PIN 3907/1) partial skull, axis (17 mm), fourth cervical vertebra, sixth cervical vertebra, eighth cervical vertebra, ninth cervical vertebra (24 mm), tenth cervical vertebra, eleventh cervical vertebra, first dorsal vertebra (~22 mm), second dorsal vertebra (~23 mm), third dorsal vertebra (~24 mm), fourth dorsal vertebra (~21 mm), fifth dorsal vertebra (~24 mm), sixth dorsal vertebra (~24 mm), seventh dorsal vertebra (~24 mm), eighth dorsal vertebra (~23 mm), ninth dorsal vertebra (~24 mm), tenth dorsal vertebra (~24 mm), two sacral ribs, partial scapulocoracoid, humerus (95 mm), proximal ulna, proximal carpometacarpus, partial ilia, pubes (~166 mm), partial ischium, femur (188 mm), tibiotarsus (257 mm), fibula, tarsometatarsus (II 143 mm, III 153 mm, IV 144 mm, V 28 mm), phalanx II-1 (~19 mm), phalanx II-2 (~13 mm), pedal ungual II (~27 mm), phalanx III-1 (~30 mm), phalanx III-2 (~21 mm), phalanx III-3 (~16 mm), pedal ungual III (~29 mm), phalanx IV-1 (~14 mm), phalanx IV-2 (~9 mm), phalanx IV-3 (~8 mm), phalanx IV-4 (~7 mm), pedal ungual IV (~28 mm)
Paratype- (PIN 3906/1) postcranial fragments including ilial fragment, proximal ischium
Referred- (PIN 3907/2) ilial fragments, proximal pubis, proximal ischium (Kurzanov, 1983)
(PIN 3907/3) anterior premaxillae, posterior skull, (mandible ~85 mm) anterior dentary, posterior mandible, skeleton including axis, tenth cervical vertebra (Kurzanov, 1985)
(PIN 3907/4) fragmentary skeleton including axis, third cervical vertebrae, fourth cervical vertebra, fifth cervical vertebra, sixth cervical vertebra, fourth dorsal vertebra (Kurzanov, 1987)
(PIN 3907/5) sacrum, ilium (~185 mm), proximal pubis, proximal ischium (Kurzanov, 1987)
(PIN 3907/6) vertebrae including sacrum (Kurzanov, 1987)
(000810 YK Avimimus UJ-1) skeleton (Watabe and Tsogtbaatar, 2004)
(000810 YK Avimimus UJ-2) skeleton (Watabe and Tsogtbaatar, 2004)
(000810 YK WTB A-IV) fragment (Tsogtbaatar and Chinzorig, 2010)
(070624 YK SZK) eight specimens (Saneyoshi, Watabe, Tsubamoto, Tsogtbaatar, Chinzorig and Suzuki, 2010)
(070628 YK ORKH) tibia (Saneyoshi, Watabe, Tsubamoto, Tsogtbaatar, Chinzorig and Suzuki, 2010)
Early Maastrichtian, Late Cretaceous
Shar Tsav, Nemegt Formation, Mongolia
(IGM 100/129; = IGM 100/120; 960822 ShT ENK) premaxillae, posterior skull, ten postaxial cervical vertebrae, about ten dorsal vertebrae, dorsal ribs, synsacrum, articulated caudal vertebrae, scapulocoracoid, sternum, humeri, radiui, ulnae, scapholunare, ulnare, carpometacarpi, phalanx II-2, partial ilia, incomplete pubes, ischium, femora, tibiotarsi, fibulae, tarsometatarsi, phalanges II-1, phalanx II-2, pedal unguals II, phalanges III-1, phalanges III-2, phalanx III-3, pedal unguals III, phalanges IV-1, phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal unguals IV (Watabe and Suzuki, 2000)
(IGM coll.; 960822 ShT SZK) pelvis, hindlimb elements (Watabe and Suzuki, 2000)
(IGM coll.; 010720 ShT-West Avimimus Ichin.) femur (Watabe, Tsogtbaatar, Ichinnorov and Barsbold, 2004)
(IGM coll.; 040804 ShT-W SZK) limb elements (Suzuki, Watabe and Tsogtbaatar, 2010)
(IGM coll.; 040804 ShT-W Tsui) (Suzuki, Watabe and Tsogtbaatar, 2010)
Diagnosis- (after Kurzanov, 1981) ilium, ischium and pubis fused around acetabulum; pelvis is wide, postacetabular processes directed almost horizontally; phalanges of pedal digit IV are highly shortened.
(after Funston et al., 2018) relatively large foramen magnum; narrower pterygoids; ilium with antitrochanter; ilium with anteriorly unexpanded preacetabular blade; postacetabular blade with suprabrevis ridges; deep brevis fossa on postacetabular blade of ilium.
(suggested) first pedal digit absent (unknown in A. nemegtensis).
Comments- Watabe et al. (2006) note Kurzanov (1981) was probably mistaken regarding the type locality of Avimimus, with Yagaan Khovil in the Nemegt Formation being more probable than Udan Sayr in the Djadockta Formation. Tsuihiji et al. (2017) incorrectly states the type locality is Shar Tsav.
Kurzanov (1981) reported two specimens of his new taxon Avimimus and described the holotype. The holotype was discovered in Summer 1973. He went on to describe the forelimb of the holotype (1982), the pelvis of the holotype and PIN 3907/2 (1983), and the skull of PIN 3907/3 (1985). His 1987 monograph described these specimens and additional ones (PIN 3907/4, 3907/5 and 3907/6) in detail, though it has yet to be translated from Russian.
Watabe et al. (2000) describe a nearly complete skeleton found in August 14 1996 that corroborates Kurzanov's identification of several features (large narial fossa in the premaxilla; narrow scapula and large coracoid; very sharp and thin posterior ulnar ridge; completely fused carpometacarpus) in addition to revealing new anatomical information (small premaxillary teeth; anteroposteriorly elongate caudals with no evidence for a pygostyle; very narrow rod-like radius). This was first noted by Watabe and Suzuki (2000) as field number 960822 ShT ENK from Shar Tsav and is photographed on page 67 as "Nearly complete skeleton of Avimimus, western area of Shar Tsav." It was later photographed after preparation by Tsogtbaatar (2004; Plate 1 Figure 6) and Matsumoto et al. (2010: Figure 1b). Tsuihiji et al. (2017) describe the premaxilla and list the specimen as IGM 100/120, but it is called IGM 100/129 by Funston et al. (2019). Found at the same time, field number 960822 ShT SZK is also from Shar Tsav and was listed by Watabe and Suzuki and Matsumoto et al. as well.
Watabe and Tsogtbaatar (2004) report two Avimimus skeletons found on August 10 2000 from Yagaan Khovil (field numbers 000810 YK Avimimus UJ-1 and UJ-2) as "cranial and postcranial elements of at least 2 individuals of Avimimus, bird-like forms including premaxilla with teeth, ilium, femur, tibia, cervical, anterior and posterior vertebrae, caudal vertebrae, digits and claws of pes (incluiding metatarsals), scapulocoracoid, ulna, carpometacarpal" and photograph one (Plate 5, Figure 1) including a dorsal vertebra, two centra, a proximal metatarsal II/IV, two pedal unguals and fragments. Tsogtbaatar (2004) lists 000810 YK as "complete pelvic, femur, scapula, vertebrae, ribs, digit", prepared in 2001. Tsogtbaatar and Chinzorig (2010) list 000810 YK Avimimus UJ-1,2 as prepared and consisting of a scapula, coracoid and pelvis, and 000810 YK WTB A-IV as consisting of a fragment.
Saneyoshi et al. (2010) reported nine additional specimens from Tagaan Khovil found in 2007.
Tsogtbaatar and Chinzorig (2010) listed a prepared femur from Shar Tsav found on July 20 2001, which was previously mentioned by Watabe et al. (2004).
References- Kurzanov, 1981. On the unusual theropods from Upper Cretaceous of Mongolia. In Resetov (ed.). Iskopaemye pozvonocnye Mongolii. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 15, 39-50.
Kurzanov, 1982. [Peculiarities of the structure of the anterior extremities of Avimimus] [in Russian]. Paleontologicheskii zhurnal. 24, 108-112.
Kurzanov, 1982. Structural characteristics of the fore limbs of Avimimus. Paleontological Journal. 16, 108-112.
Kurzanov, 1983. Avimimus and the problem of the origin of birds [in Russian]. In Resetov (ed.). Iskopaemye reptilii Mongolii. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 24, 104-109.
Kurzanov, 1983. [New data on the structure of the pelvis of Avimimus.] [in Russian]. Paleontologicheskii zhurnal. 4, 115-116.
Kurzanov, 1984. New data on the pelvic structure of Avimimus. Paleontological Journal. 17, 110-111.
Kurzanov, 1985. [The skull structure of the dinosaur Avimimus.] [in Russian]. Paleontologicheskii zhurnal. 1985, 81-89.
Kurzanov, 1985. The skull structure of the dinosaur Avimimus. Paleontological Journal. 19, 92-99.
Kurzanov, 1985. [The osteology of Avimimus portentosus and the problem of the origin of birds.] [in Russian]. Akademiya Nauk SSSR, Paleontologicheskiy Institut, Moscow. 23 pp.
Kurzanov, 1987. Avimimidae and the problem of the origin of birds [in Russian]. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 31, 1-95.
Norman, 1990. Problematic Theropoda: "Coelurosaurs". 280-305. in Weishampel, et al. (eds.). The Dinosauria. University of California Press, Berkeley, Los Angeles, Oxford.
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria (Dinosauria: Theropoda). M.S. thesis, Univ. Copenhagen, 311pp.
Dyke and Thorley, 1998. Reduced cladistic consensus methods and the avian affinities of Protoavis and Avimimus. Archaeopteryx. 16, 123-129.
Watabe and Suzuki, 2000. Report on the Japan-Mongolia Joint Paleontological Expedition to the Gobi desert, 1996. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 58-68.
Watabe, Weishampel, Barsbold, Tsogtbaatar and Suzuke, 2000. New nearly complete skeleton of the bird-like theropod, Avimimus, from the Upper Cretaceous of the Gobi Desert, Mongolia. Journal of Vertebrate Paleontology. 20(3), 77A.
Vickers-Rich, Chiappe and Kurzanov, 2002. The enigmatic birdlike dinosaur Avimimus portentosus: Comments and a pictorial atlas. In Chiappe and Witmer (eds.). Mesozoic Birds: Above the Heads of Dinosaurs. 65-86.
Tsogtbaatar, 2004. Fossil specimens prepared in Mongolian Paleontological Center 1993-2001. Hayashibara Museum of Natural Sciences Research Bulletin. 2, 123-128.
Watabe and Tsogtbaatar, 2004. Report on the Japan - Mongolia Joint Paleontological Expedition to the Gobi desert, 2000. Hayashibara Museum of Natural Sciences Research Bulletin. 2, 45-67.
Watabe, Tsogtbaatar, Ichinnorov and Barsbold, 2004. Report on the Japan - Mongolia Joint Paleontological Expedition to the Gobi desert, 2001. Hayashibara Museum of Natural Sciences Research Bulletin. 2, 69-96.
Watabe, Suzuki and Tsogtbaatar, 2006. Geological and geographical distribution of bird-like theropod, Avimimus in Mongolia. Journal of Vertebrate Paleontology. 26(3), 136A-137A.
Tsuihiji, Watabe, Barsbold, Suzuki and Tsogtbaatar, 2008. New information on the pectoral girdle, forelimb, and sternum of Avimimus portentosus (Dinosauria, Theropoda). Palaeontological Society of Japan 2008 Annual Meeting, Abstracts. 22.
Tsuihiji, Watabe, Gishlick, Barsbold and Tsogtbaatar, 2009. New information on the pectoral girdle and forelimb of Avimimus (Dinosauria: Theropoda) from the Gobi Desert of Mongolia. Journal of Vertebrate Paleontology. 29(3), 192A.
Matsumoto, Hashimoto, Sonoda, Fujiyama, Mifune, Kawahara and Saneyoshi, 2010. Report of the preparation works for Mongolian specimens in Hayashibara Museum of Natural Sciences: 1999-2008. Hayashibara Museum of Natural Sciences Research Bulletin. 3, 167-185.
Saneyoshi, Watabe, Tsubamoto, Tsogtbaatar, Chinzorig and Suzuki, 2010. Report of the HMNSMPC Joint Paleontological Expedition in 2007. Hayashibara Museum of Natural Sciences Research Bulletin. 3, 19-28.
Suzuki, Watabe and Tsogtbaatar, 2010. Report of the HMNS-MPC Joint Paleontological Expedition in 2004. Hayashibara Museum of Natural Sciences Research Bulletin. 3, 1-9.
Tsogtbaatar and Chinzorig, 2010. Fossil specimens prepared in Mongolian Paleontological Center: 2002–2008. Hayashibara Museum of Natural Sciences Research Bulletin. 3, 155-166.
Tsuihiji, Witmer, Watabe, Barsbold, Tsogtbaatar, Suzuki and Khatanbaatar, 2017. New information on the cranial morphology of Avimimus (Theropoda: Oviraptorosauria). Journal of Vertebrate Paleontology. e1347177. DOI: 10.1080/02724634.2017.1347177
Funston, Mendonca, Currie and Barsbold, 2018 (online 2017). Oviraptorosaur anatomy, diversity and ecology in the Nemegt Basin. Palaeogeography, Palaeoclimatology, Palaeoecology. 494, 101-120.
Funston, Currie, Ryan and Dong, 2019. Birdlike growth and mixed-age flocks in avimimids (Theropoda, Oviraptorosauria). Scientific Reports. 9:18816.
A. nemegtensis Funston, Mendonca, Currie and Barsbold, 2018
= Avimimus "nemegtensis" Funston, Mendonca, Currie and Barsbold, 2017 online
Early Maastrichtian, Late Cretaceous
Nemegt, Nemegt Formation, Mongolia
Holotype- (IGM 102/81) posterior skull (Funston et al., 2016)
Paratypes (IGM 102/15) proximal tibia (~254 mm)
(IGM 102/16) fused dentaries (one incomplete)
(IGM 102/17) distal tibiotarsus (~280 mm)
(IGM 102/18) distal tibiotarsus (~280 mm)
(IGM 102/19) distal tibiotarsus (tibia ~266 mm)
(IGM 102.20) distal tibiotarsus
(IGM 102/21) metatarsal II, metatarsal IV
(IGM 102/22) distal tibiotarsus (tibia ~266 mm)
(IGM 102/23) distal tibiotarsus (tibia ~265 mm)
(IGM 102/24) distal tibia (~238 mm), astragalus
(IGM 102/25) distal tibiotarsus (tibia ~265 mm)
(IGM 102/26) distal tibia (~236 mm), fibula, astragalus, calcaneum
(IGM 102/27) distal femur, proximal tibia (~232 mm)
(IGM 102/28) proximal tarsometatarsus
(IGM 102/29) proximal tarsometatarsus
(IGM 102/30) proximal tarsometatarsus
(IGM 102/31) proximal tarsometatarsus
(IGM 102/32) proximal tarsometatarsus
(IGM 102/33) (?)cranial fragment
(IGM 102/34) premaxillae, nasal
(IGM 102/35) metatarsal II
(IGM 102/36) femur
(IGM 102/37) tarsometatarsus minus metatatasal III
(IGM 102/38) (juvenile?) incomplete tibia (~202 mm)
(IGM 102/39) metatarsal II (~123 mm)
(IGM 102/40) metatarsal IV
(IGM 102/41) metatarsal IV
(IGM 102/42) tibiotarsus (269 mm; tibia 264 mm), fragments
(IGM 102/43) metatarsal
(IGM 102/44) proximal tibia (~235 mm)
(IGM 102/45) metatarsal III
(IGM 102/46) proximal tarsometatarsus
(IGM 102/47) distal tibiotarsus (tibia ~239 mm)
(IGM 102/48) metatarsal II or IV
(IGM 102/49) pubis
(IGM 102/50) distal pubes
(IGM 102/51) partial tibia (~244 mm)
(IGM 102/52) distal tibiotarsus (tibia ~246 mm)
(IGM 102/53) proximal tibia (~267 mm)
(IGM 102/54) proximal tarsometatarsus
(IGM 102/55) proximal humerus
(IGM 102/56) proximal tarsometatarsus
(IGM 102/57) proximal (?)tarsometatarsus
(IGM 102/58) proximal tarsometatarsal fragment
(IGM 102/59) proximal tarsometatarsus
(IGM 102/60) metatarsal III shaft
(IGM 102/61) distal metatarsal III
(IGM 102/62) proximal tibia (~254 mm)
(IGM 102/63) proximal tibia (~245 mm)
(IGM 102/64) fused nasals
(IGM 102/65) distal tibiotarsus
(IGM 102/66; mislabeled 102.86 in figure 7P) distal tibia (~242 mm)
(IGM 102/67) distal tibia (~231 mm)
(IGM 102/68) distal tibia (~240 mm)
(IGM 102/69) tibia (~228 mm) or pedal phalanx IV-4
(IGM 102/70) distal pubes
(IGM 102/71) metatarsal III
(IGM 102/72) proximal tibia (~273 mm)
(IGM 102/73) pubic shaft
(IGM 102/74) tibia (232 mm), fragments
(IGM 102/74A) distal tibia (~232 mm)
(IGM 102/75) proximal metatarsal
(IGM 102/76) tarsometatarsus minus metatatasal III
(IGM 102/77) metatarsal II or IV
(IGM 102/78) metatarsal II or IV
(IGM 102/79) distal metatarsal IV
(IGM 102/80) fused metatarsals
(IGM 102/82) metatarsal III
(IGM 102/83) tibia (242 mm)
(IGM 102/84) tibiotarsus (262 mm; tibia 260 mm)
(IGM 102/85) metatarsal
(IGM 102/86) two metatarsals
(IGM 102/87) metatarsal
(IGM 102/88) metatarsal
(IGM 102/89) tarsometatarsus
(IGM 102/90) tibiotarsus (264 mm; tibia 259 mm)
(IGM 102/91) metatarsal III
(IGM 102/92) tibiotarsus (282 mm; tibia 278 mm), fibula
(IGM 102/93) tarsometatarsus minus metatatasal III
(IGM 102/94) tibiotarsus (262 mm; tibia 260 mm)
(IGM 102/95) metatarsal
(IGM 102/96) tarsometatarsus minus metatatasal III
(IGM 102/97) metatarsal
(IGM 102/98) metatarsal
(IGM 102/99) metatarsal
Referred- (IGM 102/101) cervical vertebra (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(IGM 102/102) cervical verebra or tibiotarsus (tibia ~280 mm) (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(IGM 102/103) cervical vertebra (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(IGM 102/104) cervical neural arch (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(IGM 102/105) tibia (230 mm) (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(IGM 102/106) metatarsal II or IV (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(IGM 102/108) premaxillae (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.004) pedal phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.005) pedal phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.011) phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.017) vertebra (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.018) phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.022) proximal rib (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.023) rib (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.038) distal femur (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.039) proximal femur (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.040) distal femur (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.041) distal femur (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.042) pedal phalanx IV-1 (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.043) pedal phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.044) cervical vertebra (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.045) vertebra (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.048) phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.050) phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.051) fibular shaft (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.061) limb bone fragments (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.062) pedal phalanx II-1 (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.063) distal metatarsal (?)II (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.064) (?)pedal phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.067) pedal phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.068) pedal phalanx (?)III-3 (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.070) manual phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.072) proximal metatarsal II (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.073) sacral centrum (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.074) distal metatarsal (?)II (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.075) distal metatarsal III (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.076) manual phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.077) distal metatarsals IV (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.078) proximal pedal phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.079) phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.080) pedal ungual III (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.081) (?)radius (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.082) pedal phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.083) distal femur (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.085) proximal femur (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.086) proximal tibia (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.089) (?)radius (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.090) pedal ungual (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.091) pedal phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.092) tarsal (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.093) rib (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.106) ungual (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.107) rib (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.108) distal metatarsal (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.109) pedal phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.110) pedal phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.111) distal phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.112) vertebra (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.119) fibula (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.120) phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.123) rib (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.124) proximal rib (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.130) ischium (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.131) ilium (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.166) fibula (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.167) fibula (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2006.169) forelimb (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2007.023) pedal phalanx, elements (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2007.061) phalanx (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2007.062) rib (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2007.063) centrum (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2007.064) caudal vertebra (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2007.068) limb element fragment (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2007.090) fibula (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2007.092) femur (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2007.100) fibula (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2007.101) fibula (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2007.103) pedal ungual (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2007.109) vertebral fragments (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2007.111) tibia (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(PJC2007.116) fibula (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
(MPC-NEE.2016-257) scapulocoracoid (Funston, Mendonca, Currie and Barsbold, 2018)
?(ZPAL MgD-I/85) proximal tarsometatarsus (Osmólska, 1981)
eight distal caudal centra (Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016)
Early Maastrichtian, Late Cretaceous
Altan Uul, Nemegt Formation, Mongolia
?(uncollected?) material (Watabe and Suzuki, 2000)
Early Maastrichtian, Late Cretaceous
Bugin Tsav, Nemegt Formation, Mongolia
(IGM 100/125; 06816 Bgt KHTB Avi) fused nasals, jugal, posterior skull, fused dentaries, two ?cranial elements, postcrania including cervical vertebrae, rib, sacrum, caudal vertebrae, scapula, sternal, carpometacarpus, pelvis, femur and tarsometatarsus (Tsuihiji, Witmer, Watabe, Barsbold and Tsogtbaatar, 2008)
Diagnosis- (after Funston et al., 2018) exoccipital with single jugular opening; relatively small foramen magnum; wider pterygoids; laterally bowed quadratojugal; dentaries with greater development of symphyseal shelf and shallow lingual ridges; sigmoidal brevis shelf.
Comments- The description of A. nemegtensis was online November 2 2017 but only physically published April 1 2018. The type bonebed was discovered in 2006, with the holotype skull itself found on August 21 2007. At least 18 individuals are present. The nasals are incorrectly called IGM 102/46 in the main text, but are actually 102/64 with the collection number PJC2006.46 (Funston et al., 2016 supp. info).
Collected in June 8 2006, Watabe et al. (2010) first announced what would be catalogued as IGM 100/125. Although they reference "several partial skeletons of Avimimus", Appendix 1 indicates all 7 are the same specimen. Matsumoto et al. (2010) list several prepared portions. Tsuihiji et al. (2017) described a skull and identified two elements as a maxilla and lacrimal, but using the scale bars both are too large (especially the maxilla). Tsuihiji (pers. comm. 8-31-2017) confirms the scale bars are correct and at least the maxilla is too large, and notes no other individuals were present in the specimen so element misidentification is likely.
A proximal tarsometatarsus was described as Theropoda indet. by Osmólska (1981), and later identified as Avimimus (Osmólska pers. comm. to Currie, 1987; in Currie, 1989). As it is from the Nemegt type locality, it is more probably A. nemegtensis than A. portentosus.
Watabe and Suzuki (2000) reported an "unknown small theropod (possibly Avimimus)" from Altan Uul II recovered in 1997.
References- Osmólska, 1981. Coossified tarsometatarsi in theropod dinosaurs and their bearing on the problem of bird origins. Palaeontologia Polonica. 42, 79-95.
Currie, 1989. The first records of Elmisaurus (Saurischia, Theropoda) from North America. Canadian Journal of Earth Sciences. 26, 1319-1324.
Watabe and Suzuki, 2000. Report on the Japan - Mongolia Joint Paleontological Expedition to the Gobi desert, 1997. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 69-82.
Suzuki, Watabe, Saneyoshi and Tsogtbaatar, 2007. A new specimen of Avimimus (Theropoda; Oviraptorosauria) from the Upper Cretaceous Nemegt Formation and its geological age. Palaeontological Society of Japan 156th Regular Meeting, Abstracts. 17.
Currie, Longrich, Ryan, Eberth and Demchig, 2008. A bonebed of Avimimus sp. (Dinosauria: Theropoda) from the Late Cretaceous Nemegt Formation, Gobi Desert: Insights into social behavior and development in a maniraptoran theropod. Journal of Vertebrate Paleontology. 28(3), 67A.
Tsuihiji, Witmer, Watabe, Barsbold and Tsogtbaatar, 2008. New information on the cranial anatomy of Avimimus portentosus (Dinosauria: Theropoda) including virtual endocasts of the brain and inner ear. Journal of Vertebrate Paleontology. 28(3), 153A.
Matsumoto, Hashimoto, Sonoda, Fujiyama, Mifune, Kawahara and Saneyoshi, 2010. Report of the preparation works for Mongolian specimens in Hayashibara Museum of Natural Sciences: 1999-2008. Hayashibara Museum of Natural Sciences Research Bulletin. 3, 167-185.
Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010. Report of the HMNS-MPC Joint Paleontological Expedition in 2006. Hayashibara Museum of Natural Sciences Research Bulletin. 3, 11-18.
Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016. The first oviraptorosaur (Dinosauria: Theropoda) bonebed: Evidence of gregarious behaviour in a maniraptoran theropod. Scientific Reports. 6:35782. DOI: 10.1038/srep35782
Tsuihiji, Witmer, Watabe, Barsbold, Tsogtbaatar, Suzuki and Khatanbaatar, 2017. New information on the cranial morphology of Avimimus (Theropoda: Oviraptorosauria). Journal of Vertebrate Paleontology. e1347177. DOI: 10.1080/02724634.2017.1347177
Funston, Mendonca, Currie and Barsbold, 2018 (online 2017). Oviraptorosaur anatomy, diversity and ecology in the Nemegt Basin. Palaeogeography, Palaeoclimatology, Palaeoecology. 494, 101-120.
A. sp. nov. (Makovicky, 1995)
Middle-Late Campanian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China
Material- (AMNH 6570 in part; paratype of Ornithomimus asiaticus) fibula (Chiappe, Norell and Clark, 2002)
(AMNH 6576 in part; paratype of Ornithomimus asiaticus) proximal caudal vertebra (Makovicky, 1995)
?(AMNH 6754) distal metatarsal III (Ryan, Currie and Russell, 2001)
?..?(AMNH 6755; 'AMNH 6555' of Funston, Currie, Ryan and Dong, 2019) incomplete metatarsus (mtII 100, mtIV 99 mm) (Ryan, Currie and Russell, 2001)
(AMNH 25569) caudal vertebra, ten vertebrae, three phalanges, five unguals including pedal ungual (~18 mm) (AMNH online)
(AMNH coll.) dorsal vertebrae (Makovicky, 1995)
?(IVPP V16313.a) manual ungual ?I (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16314; = TMP 1992.302.0102) proximal tarsometatarsus (Ryan, Currie and Russell, 2001)
?...(IVPP V16341) tarsometatarsus (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16315) metatarsal II (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16316.a) pedal ungual III (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16316.b) pedal ungual III (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16316.c; = TMP 1992.302.0119A) pedal ungual II/IV (Ryan, Currie and Russell, 2001)
(IVPP V16316.d; = TMP 1992.302.0119B) pedal ungual II/IV (Ryan, Currie and Russell, 2001)
(IVPP V16317.a) incomplete proximal caudal vertebra (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16317.b) mid caudal vertebra (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16318; ?= TMP 1992.302.0344) (juvenile) posterior cervical vertebra (Ryan, Currie and Russell, 2001)
(IVPP V16318.a) incomplete anterior dorsal vertebra (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16318.b; ?= IVPP 160788-124) incomplete posterior dorsal vertebra (Makovicky, 1995; Funston, Currie, Ryan and Dong, 2019)
(IVPP V16319) some of "IVPP V163.." below (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16320) (<1 year old juvenile) distal tibiotarsus (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16321) proximal metatarsal II (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16322.a; = TMP 1992.302.0150) proximal tibia (Ryan, Currie and Russell, 2001)
(IVPP V16322.b) proximal tibia (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16322.c) proximal tibia (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16323.a) third dorsal vertebra (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16324) some of "IVPP V163.." below (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16325) some of "IVPP V163.." below (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16326) metatarsal II (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16327; = TMP 1992.302.0116) partial scapulocoracoid (Ryan, Currie and Russell, 2001)
(IVPP V16328) (juvenile) mid sacral centrum (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16329.a; = IVPP 180788-123) last or penultimate cervical vertebra (Makovicky, 1995)
(IVPP V16329.b) second dorsal vertebra (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16330; = IVPP 160788-122) posterior sacral fragment (Makovicky, 1995)
(IVPP V16331) some of "IVPP V163.." below (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16332.a; = TMP 1992.302.0140) partial second dorsal vertebra (Ryan, Currie and Russell, 2001)
(IVPP V16332.b) partial first dorsal vertebra (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16333) some of "IVPP V163.." below (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16334.a; = TMP 1992.302.0149) proximal femur (Ryan, Currie and Russell, 2001)
(IVPP V16335.a) (juvenile) distal tarsal IV fused to proximal metatarsal IV (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16335.b) distal metatarsal II/III
....(IVPP V16335.c) distal metatarsal II/III
....(IVPP V16336) distal metatarsal IV
(IVPP V16337) (>2 year old adult) distal tarsometatarsus (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16338; = TMP 1992.302.0110) distal femur (Ryan, Currie and Russell, 2001)
(IVPP V16339) some of "IVPP V163.." below (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16340; = TMP 1992.302.0117) proximal humerus (Ryan, Currie and Russell, 2001)
?(IVPP V16342; = TMP 1992.302.0104) partial frontal (Ryan, Currie and Russell, 2001)
(IVPP V16343) distal metacarpal I (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16344) some of "IVPP V163.." below (Funston, Currie, Ryan and Dong, 2019)
(IVPP V16345) some of "IVPP V163.." below (Funston, Currie, Ryan and Dong, 2019)
(IVPP V163...) four dorsal vertebrae, distal metatarsal, pedal ungual III, eight pedal unguals II/IV (Makovicky, 1995; Funston, Currie, Ryan and Dong, 2019)
(PIN coll.) material (Currie and Eberth, 1993)
Diagnosis- (after Funston et al., 2019) three cervicodorsal vertebrae (unknown in A. nemegtensis); deeper femoral intercondylar groove.
Comments- Makovicky (1995) stated "Avimimid vertebrae were collected by the American Museum Central Asian expedition in 1922 at Iren Dabasu. However, they were not recognised as such, and were catalogued with Ornithomimus asiaticus (now Archaeornithomimus asiaticus) material", but the Archaeornithomimus type material was discovered in 1923 including a proximal caudal referenced by the author. While he says "one of the caudals collected by the American Museum's Central Asiatic Expeditions (AMNH 6576), [has] a small pointed tubercle separates the ventromedial corners of the chevron facets" this specimen number includes almost a hundred elements from the Johnson Quarry AMNH locality 141 generally referred to Archaeornithomimus. A fibula (AMNH 6570) was figured as "alvarezsaurid fibula from Iren Dabasu (Inner Mongolia, China)" by Chiappe et al. (2002), but Longrich and Currie (2009) stated "it more closely resembles the fibula of Avimimidae, which are common at this locality (N.R.L., pers. obs.)." Again this number includes over two hundred paratype Archaeornithomimus elements, this time from the Kaisen Quarry AMNH locality 140. Makovicky also stated "A large number of dorsal vertebrae are present in the American Museum ... collections from Iren Dabasu", which are likely to also be catalogued under AMNH 6570 and/or 6576. Longrich's claim is supported here however as the fibula has an anteriorly projected iliofibularis tuber as in Avimimus but unlike the laterally directed tuber of e.g. IGM 100/99 and is less reduced distally in anteroposterior width than the latter. A distal caudal (AMNH coll.) described by Makovicky "has the morphology of a typical coelurosaurian distal caudal, but is otherwise undiagnostic. The possibility that it may originate from an avimimid is suggested by its small size. It should be noted, however, that it could just as conceivably be from the tail of a juvenile Archaeornithomimus from the same bonebed." Indeed, the A. nemegtensis bonebed show Avimimus has short distal caudals like other caenagnathoids and unlike ornithomimosaurs, so this specimen is here referred to Archaeornithomimus. The AMNH online catalogue lists AMNH 25569 as "10 vertebrae 1 caudal vertebra 3 phalanges & 5 claws" of Saurischia from the Johnson Quarry with a pedal ungual photographed as Avimimidae. A metatarsus (AMNH 6755) and third metatarsal (AMNH 6754) were listed on the museum's online catalogue as Elmisaurus sp., but also referred to Avimimus by Ryan et al. (2001) and Funston et al. (for AMNH 6755 at least). Ryan et al.'s poster indicated both were found in 1923 and were possibly associated. AMNH 6755 does seem smaller than other fused Avimimus and less slender, so further study is necessary. Note Funston et al. (2019) describe and figure this as AMNH 6555, which is the number of ornithischian material. Funston et al. state the main Iren Dabasu Avimimus bonebed "was originally discovered by a Sino-Soviet expedition in 1959, which used bulldozers to excavate the site. It was revisited in 1987 and 1988 by the Sino-Canadian expedition and numerous fragmentary bones representing all regions of the skeleton (Fig. 2) were recovered from the spoil piles left by the Sino-Soviet bulldozers. Unfortunately, the material collected by the Sino-Soviet expedition still awaits preparation and it may never be available for study." This indicates it must be locality K of Currie and Eberth (1993), which was CCDP locality 1 and may correspond to AMNH locality 141. One element from this excavation was described and figured by Kurzanov (1987) as "a left avimimid femur from the Upper Cretaceous Iren-Nor locality in China (specimen PIN, no. 2549-100)" (translated) but is here placed in Oviraptoridae. While the PIN Sino-Soviet Avimimus material remains undescribed (although mentioned by Currie and Eberth), the Sino-Canadian material was noted by Dong et al. (1989) who reported "seven pedal elements of Avimimus" discovered in July 1988 and Dong (1992) refers to "fused tarsometatarsi of Avimimus" recovered on that expedition in July 1998. Currie and Eberth (1993) stated "Direct comparison between isolated Avimimus bones from the Iren Dabasu and the type specimen of Avimimus portentosus in Moscow failed to reveal any differences" and said the "material is presently under review (Currie, Zhao and Kurzanov, in preparation)." The Sino-Canadian Avimimus material was eventually described by Makovicky, Ryan et al., then officially by Funston et al. as Avimimus sp.. Makovicky used IVPP field numbers (of which IVPP 180788-123 may be a mistake for IVPP 160788-123 to better match the other two listed numbers and only differ in the last digit), Ryan et al. used TMP numbers in their poster, and the specimens were seemingly eventually transported back to the IVPP for permanent storage. Note the "isolated left frontal" mentioned by Sues et al. is not described in Funston et al. (although the frontal is highlighted as preserved in their figure 2), but Funston (2019) describes it in his thesis chapter that was developed into that paper. In it he says "whether it pertains to an avimimid or another oviraptorosaur is uncertain. It can be distinguished from other theropods by the large, incising nasal contact, which is similar to the morphology of Elmisaurus rarus", so it's possible this belonged to the Iren Dabasu caenagnathid taxon represented by dentaries IVPP V20377. IVPP 160788-124 is one of three posterior dorsals without a ventral keel, so may be IVPP V16318.b which was the only one of these figured by Funston et al.. Another tricky specimen is TMP 1992.302.0344 as illustrated in Ryan et al.'s poster, which is tentatively identified as posterior cervical IVPP V16318 as it seems to have a ventrally placed parapophysis, an elongate centrum with vertical articular surfaces, two or three central foramina and a dark neurocentral boundary perhaps representing an open suture, but contra to Funston et al.'s description is not notably smaller than other cervicals. While Funston et al. only referred to the Iren Dabasu material as Avimimidae gen. et sp. indet., they also stated "the cervicodorsal vertebrae differ in number (three with hypapophyses) from those of Avimimus portentosus (MPC-D 100/129), although cervicodorsal number in Avimimus nemegtensis is unknown. The distal condyles of the femur (Fig. 6) are separated much more deeply than is typical in avimimids, and metatarsals II and IV (Fig. 8) are much more disparate in size", but ended up concluding "the available material from the Iren Dabasu bonebed is too incomplete to confidently erect a new taxon, but future preparation of the Russian material (or collection of new material) may result in its taxonomic distinction from other avimimids."
References- Kurzanov, 1987. Avimimidae and the problem of the origin of birds. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 31, 1-95.
Dong, Currie and Russell, 1989. The 1988 field program of the Dinosaur Project. Vertebrata Palasiatica. 27(3), 233-236.
Dong, 1992. Dinosaurian Faunas of China. China Ocean Press. 188 pp.
Currie and Eberth, 1993. Palaeontology, sedimentology and palaeoecology of the Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia, People's Republic of China. Cretaceous Research. 14, 127-144.
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria (Dinosauria: Theropoda). Masters thesis, University of Copenhagen. 311 pp.
Ryan, Currie and Russell, 2001. New material of Avimimus portentosus (Theropoda) from the Iren Dabasu Formation (Upper Cretaceous) of the Erenhot region of Inner Mongolia. Journal of Vertebrate Paleontology. 21(3), 95A.
Chiappe, Norell and Clark, 2002. The Cretaceous, short-armed Alvarezsauridae, Mononykus and its kin. In Chiappe and Witmer (eds.). Mesozoic Birds: Above the Heads of Dinosaurs. University of California Press. 87-120.
Longrich and Currie, 2009 (online 2008). Albertonykus borealis, a new alvarezsaur (Dinosauria: Theropoda) from the Early Maastrichtian of Alberta, Canada: Implications for the systematics and ecology of the Alvarezsauridae. Cretaceous Research. 30(1), 239-252.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria (Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Funston, Currie, Ryan and Dong, 2019. Birdlike growth and mixed-age flocks in avimimids (Theropoda, Oviraptorosauria). Scientific Reports. 9:18816.
A. sp. (Currie, 2000)
Cenomanian-Santonian, Late Cretaceous
Bayshin Tsav, Baynshiren Formation, Mongolia
Material- (IGM coll.) isolated elements
Comments- Currie (2000) stated "isolated avimimid bones have been collected ... in 1975 at Baishin Tsav (Kurzanov, pers. comm., 1991)."
Reference- Currie, 2000. Theropod dinosaurs from the Cretaceous of Mongolia. In Benton, Shishkin, Unwin and Kurochkin (eds.). The Age of Dinosaurs in Russia and Mongolia. 434-455.
A. sp. (Jerzykiewicz and Russell, 1991)
Late Campanian, Late Cretaceous
Djadokhta Formation, Mongolia
Comments- Jerzykiewicz and Russell (1991) list "Avimimus portentosus (Kurzanov, pers. comm., 1988)" under the Djadochta Formation.
Reference- Jerzykiewicz and Russell, 1991. Late Mesozoic stratigraphy and vertebrates of the Gobi Basin. Cretaceous Research. 12, 345-377.
A. sp. (Averianov and Lopatin, 2022a)
Early Santonian, Late Cretaceous
Kansai, Yalovach Formation, Tajikistan
Material- (PIN 3041/49) distal tibiotarsus (30.5 mm trans)
Comments- Discovered in 1963-1964, Averianov and Lopatin (2022b) stated that compared to the A. portentosus holotype "The Kansai specimen is distinguished by the presence of a fusiform depression on the anterior side of the tibiotarsus proximal to the distal condyles, as well as a more developed postfibular flange and a transverse ridge on the medial condyle. These differences may be related to the greater ontogenetic age of the Kansai specimen. This clear similarity allows us to identify the specimen PIN, no. 3041/49 as Avimimus sp.".
References- Averianov and Lopatin, 2022a. First discovery of theropod dinosaurs of the family Avimimidae in the Late Cretaceous of central Asia. Doklady Earth Sciences. 506(2), 210-213.
Averianov and Lopatin, 2022b. First discovery of theropod dinosaurs of the family Avimimidae in the Late Cretaceous of central Asia. Doklady Earth Sciences. 506(2), 775-777.
A. spp. (Makovicky, 1995)
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada
Material- (TMP 1980.029.0096) metatarsal (TMP online)
(TMP 1989.069.0012) metatarsal (TMP online)
(TMP 1990.151.0006) incomplete limb element (TMP online)
(TMP 1994.012.0384) pedal ungual (TMP online)
(TMP 1994.089.0017) metatarsal (TMP online)
(TMP 1998.068.0022) distal metatarsal III (Ryan, Currie and Russell, 2001)
(TMP 1998.068.0092) metatarsal (TMP online)
(TMP 1998.068.0102) phalanx (TMP online)
(TMP coll.) vertebrae (Currie, 2000)
Early Maastrichtian, Late Cretaceous
Horseshoe Canyon Formation, Alberta, Canada
(TMP or UALVP coll.) (unassociated) metatarsal III, distal metatarsal, ?pedal phalanx (Eberth and Currie, 2010)
Late Maastrichtian, Late Cretaceous
Scollard Formation, Alberta, Canada
(TMP 1998.008.0028) distal metatarsal II (Ryan and Russell, 2001)
Comments- Makovicky (1995) first noted "Isolated bones [of Avimimus] have also been found in North America, including the Dinosaur Park formation of Alberta (Currie, in prep.)." Currie (2000) stated "A number of isolated vertebrae, tarsometatarsi and unguals have been found in Upper Cretaceous strata of North America that closely resemble those of Mongolian avimimids (collections of Royal Tyrrell Museum of Palaeontology)", but Ryan et al. (2001) found that although there are twelve elements in the TMP collections labeled Avimimidae, only two can certainly be assigned to that taxon. One, TMP 1998.068.0022, is a distal third metatarsal from the Dinosaur Park Formation. The other (TMP 1998.008.0028) is a second metatarsal from the Scollard Formation that is unfused proximally, unlike A. portentosus. This was listed as cf. Avimimus sp. in Ryan and Russell (2001) and the TMP online catalogue includes a photo that indicates it only preserves the distal end. The TMP online catalogue includes nine avimimid records including metatarsi and a pedal ungual, but no vertebrae. Note TMP 1989.069.0012 and 1990.151.0006 are listed as being from the Oldman Formation, but the Onefour locality they were discovered in is in the Dinosaur Park Formation. Funston (2019) states regarding Ryan et al.'s specimens, "re-examination indicates that this material more likely belongs to another small theropod" but with no rationale. Eberth and Currie (2010) listed three avimimid elements from the Albertosaurus bonebed of the Horseshoe Canyon Formation.
References- Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria (Dinosauria: Theropoda). Masters thesis, University of Copenhagen. 311 pp.
Currie, 2000. Theropod dinosaurs from the Cretaceous of Mongolia. In Benton, Shishkin, Unwin and Kurochkin (eds.). The Age of Dinosaurs in Russia and Mongolia. 434-455.
Ryan, Currie and Russell, 2001. New material of Avimimus portentosus (Theropoda) from the Iren Dabasu Formation (Upper Cretaceous) of the Erenhot region of Inner Mongolia. Journal of Vertebrate Paleontology. 21(3), 95A.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the Paleontology of Philip J. Currie. Indiana University Press. 279-297.
Ryan and Currie, 2002. Asian small theropods in North America: Evidence from Avimimidae. In Alberta Palaeontological Society, sixth annual symposium. 44.
Currie, 2005. Theropod dinosaurs of Dinosaur Provincial Park. In Braman, Therrien, Koppelhus and Taylor (eds.). Dinosaur Park Symposium, short papers, abstracts and program, special publication of the Royal Tyrrell Museum. 15-18.
Eberth and Currie, 2010. Stratigraphy, sedimentology, and taphonomy of the Albertosaurus bonebed (upper Horseshoe Canyon Formation; Maastrichtian), southern Alberta, Canada. Canadian Journal of Earth Sciences. 47(9), 1119-1143.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria (Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.

Oviraptoridae Barsbold, 1976a
Definition- (Oviraptor philoceratops <- Caenagnathus collinsi) (Maryanska, Osmólska and Wolsan, 2002; modified from Sereno, 1998)
Other definitions- (Oviraptor philoceratops <- Chirostenotes pergracilis) (Sereno, online 2005; modified from Padian, Hutchinson and Holtz, 1999)
= Ingeniidae Barsbold, 1981
= Oviraptoridae sensu Padian, Hutchinson and Holtz, 1999
definition- (Oviraptor philoceratops <- Chirostenotes pergracilis) (modified)
Comments- Supposed oviraptorid material from the Yalovach Formation of Tadjikistan (Nessov, 1995) probably belongs to therizinosaurs instead (Alifanov and Averianov, 2006).
References- Barsbold, 1976a. O novum pozdnemelovom semeystve melkikh teropod (Oviraptoridae fem. n.) Mongolii. Doklady Akademii Nauk SSSR. 226(3), 685-688.
Barsbold, 1976b. A new Late Cretaceous family of small theropods (Oviraptoridae n. fam.) in Mongolia. Doklady Akademia Nauk SSSR. 226, 221-223.
Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological Expedition Transactions. 15, 28-39.
Nessov, 1995. Dinosaurs of northern Eurasia: New data about assemblages, ecology and paleobiogeography. Scientific Research Institute of the Earth's Crust, St. Petersburg State University, St. Petersburg, Russia. 156 pp.
Sereno, 1998. A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen. 210(1), 41-83.
Padian, Hutchinson and Holtz, 1999. Phylogenetic definitions and nomenclature of the major taxonomic categories of the carnivorous Dinosauria (Theropoda). Journal of Vertebrate Paleontology. 19(1), 69-80.
Maryanska, Osmólska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Sereno, online 2005. Stem Archosauria - TaxonSearch. http://www.taxonsearch.org/dev/file_home.php [version 1.0, 2005 November 7]
Alifanov and Averianov, 2006. On the finding of ornithomimid dinosaurs (Saurischia, Ornithomimosauria) in the Upper Cretaceous beds of Tajikistan. Paleontological Journal. 40(1), 103-108.
Smith and Molnar, 2006. Jaw musculature and function in oviraptorosaurs. Journal of Vertebrate Paleontology. 26(3), 126A.

Shanyangosaurus Xue, Zhang and Bi, 1996
S. niupanggouensis Xue, Zhang and Bi, 1996
Middle-Late Maastrichtian, Late Cretaceous
Shanyang Formation, Shaanxi, China
Holotype- (NWUV 1111) (~1.7 m) uncinate processes(?), partial synsacrum (centra 32 mm), proximal scapula, humeri (116 mm), femur (258 mm), tibia (327 mm), metatarsals IV (137 mm; one proximal), partial phalanx, pedal ungual
Comments- Xue et al. (1996) report "ribs with horizontal hooks", but no ribs are mentioned in the material list, nor are any shown in the plates. If these are uncinate processes, they would support a pennaraptoran identity.
While Shanyangosaurus was originally identified only to the level of Theropoda, it is most similar to oviraptorids (Mortimer, DML 2000; followed by Holtz et al., 2004) in the low acromion, absent fourth trochanter, cnemial crest shape, unfused metatarsus whose fourth metatarsal is wider than deep, and an elevated femoral head. Hartman et al. (2019) were the first authors to include it in a phylogenetic analysis, finding it emerged in Caudipteridae. Yet as it moves to Oviraptoridae in one step, the cnemial crest shape was not analyzed, and the age is more congruent with oviraptorids it is tentatively placed there on this site. If it is constrained as an oviraptorid in an updated version of the analysis, it falls outside the "Huanansaurus" plus Oviraptor clade.
References- Xue, Zhang, Bi, Yue and Chen, 1996. The development and environmental changes of the intermontane basins in the eastern part of Qinling Mountains. Geological Publishing House, Beijing. ISBN 7-116-02125-6. 179 pp.
Mortimer, DML 2000. https://web.archive.org/web/20210506115234/http://dml.cmnh.org/2000Sep/msg00125.html
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

undescribed possible Oviraptoridae (Kirkland, Hernandez Rivera, Aguillon Martinez, Delgado de Jesus, Gomez Nunez and Vallejo, 2000)
Campanian, Late Cretaceous
Cerro del Pueblo Formation, Mexico
Comments- Kirkland et al. (2000) stated "theropods are represented by isolated bones (Fig. 13C), claws, and teeth indicating the presence of tyrannosaurids, dromaeosaurids, ornithomimids, and oviraptorids." Figure 13C is a hadrosaur femur, so the authors apparently meant figure 12C consisting of theropod phalanges. Some of these appear to be elements from the El Pelillal locality (e.g. IGM-7710 and 7712) described by Rodriguez-De La Rosa and Cevallos-Ferriz (1998), but Aguillon Martinez (2010) noted this locality is actually from the younger Cañon del Tule Formation. Although Aguillon Martinez stated "small vertebrates from the Cerro del Pueblo Formation remain undocumented", Rodriguez-De La Rosa and Cevallos-Ferriz do indicate "Theropoda Indet. family" remains are known from several areas of the Cerro del Pueblo Formation- Presa San Antonio, Agua de la Mula, La Rosa and Rincon Colorado area. That being said, no oviraptorosaur material was noted by Aguillon Martinez or Rivera-Sylva and Carpenter (2014) and so may have been misidentified.
References- Rodriguez-De La Rosa and Cevallos-Ferriz, 1998. Vertebrates of the El Pelillal locality (Campanian, Cerro del Pueblo Formation), southeastern Coahuila, Mexico. Journal of Vertebrate Paleontology. 18(4), 751-764.
Kirkland, Hernandez-Rivera, Aguillón Martinez, de Jesus, Gomez-Nunez and Vallejo, 2000. The Late Cretaceous Difunta Group of the Parras Basin, Coahuila, Mexico and its vertebrate fauna. Universidad Autónoma del Estado de Hidalgo, Avances en Investigación. 3, 133-172.
Aguillon Martinez, 2010. Fossil vertebrates from the Cerro del Pueblo Formation, Coahuila, Mexico, and the distribution of Late Campanian (Cretaceous) terrestrial vertebrate faunas. MS thesis, Dedman College Southern Methodist University. 135 pp.
Rivera-Sylva and Carpenter, 2014. Mexican saurischian dinosaurs. In Rivera-Sylva, Carpenter and Frey (eds.). Dinosaurs and Other Reptiles from the Mesozoic of Mexico. Indiana University Press. 143-155.

unnamed possible Oviraptoridae (Riabinin, 1938)
Santonian, Late Cretaceous
Syuk Syuk Formation, Kazakhstan
Material- unguals
References- Prinada, 1925. Search for remains of large vertebrates of Upper Cretaceous age in Turkestan. Report on the state of activities of the Geological Committee for 1924. Part II, III. Izvyestiya Gyeologichyeskogo komityeta 44(2): 257.
Prinada, 1927. Report on the excavation at the localities where dinosaur bones were discovered. Report on the state of activities of the Geological Committee for 1925. Part II, III. Izvyestiya Gyeologichyeskogo komityeta 45(4): 453-454.
Riabinin, 1938. Some results of the study of the Upper Cretaceous dinosaur fauna from the vicinity of st. Sary-Agachin, Southern Kazakhstan. Problyemy palyeontologii 4: 125-135.
Nessov, 1995. Dinosaurs of nothern Eurasia: new data about assemblages, ecology, and paleobiogeography. Institute for Scientific Research on the Earth's Crust, St. Petersburg State University, St. Petersburg 1-156.

undescribed Oviraptoridae (Fanti et al., 2012)
Late Cretaceous(?)
Mongolia or China(?)
Material- (IGM coll.) specimen including skull (147.3 mm), mandible (120 mm), metacarpal I (24 mm), phalanx I-1 (30.1 mm), manual ungual I (31.7 mm), phalanx II-2 (17.8 mm), manual ungual II (81.8 mm), femur (238 mm), tibia (275 mm), phalanx III-3 (17 mm), phalanx IV-2 (13.3 mm), phalanx IV-3 (10 mm), phalanx IV-4 (9.3 mm), pedal ungual IV (30 mm) (Fanti et al., 2012)
(IGM coll.) specimen including skull (162 mm), humerus (113 mm), radius (90 mm), ulna (~98 mm), metacarpal I (22.4 mm), phalanx I-1 (29.2 mm), manual ungual I (35.2 mm), metacarpal II (40.6 mm), phalanx II-1 (19.6 mm), phalanx II-2 (15 mm), manual ungual II (18 mm), femur (220 mm), tibia (265 mm), phalanx III-3 (15.8 mm), phalanx IV-2 (13.6 mm), phalanx IV-3 (9.9 mm), phalanx IV-4 (8.9 mm), pedal ungual IV (27.3 mm) (Fanti et al., 2012)
skull, mandible, five anterior cervical vertebrae (www.dino-pantheon.com, online 2007)
Comments- Fanti et al. (2012) listed measurements for two "oviraptorid incertae sedis" specimens at the IGM. Analyzing their proportions may enable identification. Numerous unpublished oviraptorid specimens are known and have been photographed online. Most are referred to Oviraptor, Conchoraptor or "Ingenia", but this is generally based on the pre-1980's convention of calling all oviraptorids Oviraptor or the pre-2000 convention of calling all crestless oviraptorids Conchoraptor or "Ingenia". The more complete skeletons, especially when mounted, often contain faked portions.
References- www.dino-pantheon.com, online 2007. http://www.dino-pantheon.com/museumreport/photo/oviraptorhead.jpg
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.

undescribed oviraptorid (U.S. Attorney's Office Southern District of New York, online 2014)
Late Cretaceous(?)
Mongolia
Material- (CMMD coll.) skeleton
Comments- U.S. Attorney's Office Southern District of New York (online 2014) report on a legal case where smuggled dinosaurs were being returned to the CMMD in Mongolia, including "two freestanding Oviraptors." Only one has been photographed online to my knowledge, so the identity and amount of real fossil in the second mount remains unknown.
Reference- U.S. Attorney's Office Southern District of New York, online 2014. Manhattan U.S. Attorney Announces Return To Mongolia Of Fossils Of Over 18 Dinosaur Skeletons. July 10.

undescribed oviraptorid (Lee, Barsbold, Jacobs and Currie, 2008)
Cenomanian-Early Maastrichtian, Late Cretaceous
Baynshiren, Baruungoyot or Nemegt Formation, Mongolia
Material- (IGM coll.) embryos, nest
Comments- Discovered in 2007 and noted as "an oviraptorosaurid nest with embryos." Expeditions that year were from "the Bayanshiree Formation at Bayan Shiree, Khar Khutul, and Shine Us Khudag in the eastern Gobi, and the Baruungoyot and Nemegt formations at Khermeen Tsav in the western Gobi."
References- Lee and Barsbold, 2008. Korea-Mongolia International Dinosaur Project. Abstracts of the International Dinosaur Symposium in Fukui 2008. 27-28.
Lee, Barsbold, Jacobs and Currie, 2008. A short report of Korea-Mongolia International Dinosaur Project (1st and 2nd year). Journal of Vertebrate Paleontology. 28(3), 104A-105A.

undescribed Oviraptoridae (Norell, Balanoff, Barta and Erickson, 2018)
Late Campanian, Late Cretaceous
Dzamin Khond, Djadochta Formation, Mongolia
Material- (IGM 100/79-D; labeled IGM 100/79-A) skull, mandibles, atlas, axis (Funston, 2019)
Late Campanian, Late Cretaceous
Udyn Sayr, Djadokhta Formation, Mongolia
(IGM 16-08) multiple individuals (Norell, Balanoff, Barta and Erickson, 2018)
Comments- Norell et al. (2018) mention an assemblage of individuals from Udan Sayr.
On display at the Mongolian Natural History Museum in 2002 (photographed by Tweet, 2019 online), an oviraptorid skull is labeled 100/79-A. It was described in Funston's (2019) thesis as a new taxon, which calls it MPC-D 100/79D and reveals the locality to be Dzamin Khond.
References- Norell, Balanoff, Barta and Erickson, 2018. A second specimen of Citipati Osmólskae associated with a nest of eggs from Ukhaa Tolgod, Omnogov Aimag, Mongolia. American Museum Novitates. 3899, 44 pp.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria (Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Tweet, 2019 online. https://1.bp.blogspot.com/-ODfgpelZv3w/XWIGpz_6fKI/AAAAAAAADWo/j-A4E6S2FKQ80kO8q2bDVpl1GxAtvMO9QCLcBGAs/s1600/oviraptor001.jpg

undescribed oviraptorid (Jensen, 2008)
Late Campanian(?), Late Cretaceous
Red Beds of Khermeen Tsav, Baruungoyot Formation, Mongolia
Material- ?(CMMD coll.) skull, mandibles, eleven cervical vertebrae, cervical ribs, thirteen dorsal vertebrae, dorsal ribs, sacrum, caudal series, chevrons, scapulae, coracoids, furcula, humeri, radii, ulnae, semilunate carpal, metacarpals I, phalanges I-1, manual unguals I, metacarpals II, phalanges II-1, phalanges II-2, manual ungual II, metacarpals III, phalanges III-1, phalanges III-2, phalanges III-3, manual unguals III, ilium, pubes, ischia, femora, tibiae, fibulae, astragali, metatarsals II, phalanges II-1, phalanges II-2, pedal unguals II, metatarsals III, phalanges III-1, phalanges III-2, phalanges III-3, pedal unguals III, metatarsals IV, phalanges IV-1, phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal unguals IV
(private coll.; PMO X678 cast; UALVP 49394 cast) skull (112 mm), mandibles (99.8 mm), hyoids, eight partial cervical vertebrae, dorsal rib fragments (Knutsen, 2008)
(private coll.; UALVL 49393 cast) skull (~126 mm), sclerotic ring, mandibles (139 mm), hyoids, phalanx I-1 (57.5 mm), manual ungual I (47 mm), incomplete metacarpal II, phalanx II-1 (56 mm), phalanx II-2 (55 mm), manual ungual II (47 mm), incomplete metacarpal III, phalanx III-1 (38 mm), phalanx III-2, phalanx III-3, manual ungual III (Fanti et al., 2012)
Comments- This taxon is known from three specimens with similar high, pointed crests. Casts are common, with Prehistoric Planet Store (online, 2019) listing one as being from Khermeen Tsav. This specimen was analyzed by Jensen (2008), who called it Oviraptor sp..
The second specimen includes a manus preserved next to the skull and is listed as Oviraptor philoceratops on the Witmer Lab website (Witmer, 2012 online). As noted by Rey (DML, 2003), Gee and Rey (2003) based their fictional oviraptorosaur "Ronaldoraptor" on this skull, but as the taxon is not obviously nonfictional in the book it is not considered actual nomenclature here. Fanti et al. (2012) used UALVP casts (Oviraptor in their online catalogue) to include measurements of both specimens in their dataset as "oviraptorid incertae sedis".
U.S. Attorney's Office Southern District of New York (online 2014) report on a legal case where smuggled dinosaurs were being returned to the CMMD in Mongolia, including "two freestanding Oviraptors." One of these is photographed by Todd (2017 online) and has a cranial crest similar in shape to the above specimens. While at least some of the mounted skeleton must be real for it to be illegal to own (and indeed the e.g. proximal scapula, some dorsal ribs and ilium seem to be actual fossils), the texture of the forelimbs, feet otherwise complete but both lacking digit I and metatarsal V, and low cervicodorsal ratio suggest at least some is a cast. This includes the possibility the skull is cast, which would eliminate the only reason it is listed under this entry here, and it should be noted the manus strongly differs from the UALVL 49393 specimen as well, with shorter phalanges and a larger ungual I. The manus may be cast as well of course. Making things even more ambiguous, the specimen is listed as being from the Djadokhta Formation on its CMMD plaque, but the associated point on the map is far to the west in Gurvantes, close to Nemegt and Baynshiren loacilities but far from Djadokhta localities.
References- Gee and Rey, 2003. A Field Guide to Dinosaurs: The Essential Handbook for Travelers in the Mesozoic. Barron's Educational Series. 144 pp.
Rey, DML 2003. https://web.archive.org/web/20210603185800/http://dml.cmnh.org/2003Mar/msg00210.html
Jensen, 2008. Beak morphology in oviraptorids, based on extant birds and turtles. Masters Thesis. University of Oslo. 48 pp.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
Witmer, 2012 online. https://people.ohio.edu/witmerl/collections/Theropods/oviraptor.htm
U.S. Attorney's Office Southern District of New York, online 2014. Manhattan U.S. Attorney Announces Return To Mongolia Of Fossils Of Over 18 Dinosaur Skeletons. July 10.
Todd, 2017 online. https://www.flickr.com/photos/101561334@N08/35724849645/in/album-72157683092773953/
Prehistoric Planet Store online, 2019. https://web.archive.org/web/20210120030224/http://www.prehistoricstore.com/item.php?item=1839

unnamed Oviraptoridae (Maryanska and Osmólska, 1997)
Late Campanian(?), Late Cretaceous
Red Beds of Khermeen Tsav, Baruungoyot Formation, Mongolia
Material- (IGM 100/30A) fragmentary skull (quadrate 50 mm) (Maryanska and Osmólska, 1997)
(IGM coll.; "GIN A") quadrates (52 mm) (Maryanska and Osmólska, 1997)
(IGM coll.; "GIN B") partial quadratojugal, distal quadrate (Maryanska and Osmólska, 1997)
(IGM coll.) skull (Clark, 1992)
Comments- These may be Conchoraptor, Heyuannia or Nemegtomaia based on their provenence.
The skull preliminarily reported by Clark (1992) as Oviraptor sp.was found in 1991. The other material was stated by Maryanska and Osmólska (1997) to "represent uncrested oviraptorids (probably Ingenia yanshini Barsbold, 1981 or Conchoraptor gracilis Barsbold, 1986)", but how this can be determined from such fragmentary remains as GIN A or GIN B is uncertain.
References- Clark, 1992. The Mongolian-American Museum Expeditions to the Gobi Desert, 1990-1992. Journal of Vertebrate Paleontology. 12(3), 24A.
Maryanska and Osmólska, 1997. The quadrate of oviraptorid dinosaurs. Acta Palaeontologia Polonica. 42, 377-387.

unnamed Oviraptoridae (Osmólska, 1976)
Early Maastrichtian, Late Cretaceous
Bugin Tsav, Nemegt Formation, Mongolia
Material- (IGM 100/1017) (embryo) femur, fragmentary bones, partial egg (Weishampel et al., 2008)
(IGM 100/1018) (embryo) partial skull, surangular?, six cervical vertebrae, few cervical ribs, three dorsal vertebrae, dorsal ribs, scapula, coracoid, humerus (17 mm), radius, ulna, metacarpal I, metacarpal II, metacarpal III, ilia, femora (30 mm), tibia (42 mm), distal fibula, astragalus, metatarsal II, metatarsal III, pedal ungual III, metatarsal IV, several pedal phalanges, eggshell (Weishampel et al., 2008)
(IGM 100/1019-1) (embryo) partial skull, surangular or articular, three cervical vertebrae, humerus (22 mm), eggshell (Weishampel et al., 2008)
(IGM 100/1019-2) (embryo) dorsal vertebrae, ilium, femur (39 mm), tibia (55 mm), fibula, astragalus (Weishampel et al., 2008)
Early Maastrichtian, Late Cretaceous
Gurilin Tsav, Nemegt Formation, Mongolia
(IGM coll.; 980807 GT Coy Oviraptr) partial ?cervical vertebra, five dorsal vertebrae, five proximal dorsal ribs, synsacrum, ventral ilia, pubes, incomplete ischia (Suzuki and Watabe, 2000)
Early Maastrichtian, Late Cretaceous
Khaichin Uul, Nemegt Formation, Mongolia
three dorsal vertebrae, three caudal vertebrae (Badamkhatan, 2008)
Early Maastrichtian, Late Cretaceous
Khermeen Tsav, Nemegt Formation, Mongolia
partial skeleton (Currie, 2001)
Early Maastrichtian, Late Cretaceous
Nemegt, Nemegt Formation, Mongolia
elements (Currie et al., 2008)
Early Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia
(ZPAL MgD-I/96) fragmentary skull (Osmólska, 1976)
Comments- These may be Nomingia, Rinchenia, Gobiraptor or Nemegtomaia based on their provenence.
Osmólska (1976) mentioned a fragmentary skull from the Nemegt Formation of Mongolia housed in the ZPAL. She referred it to Oviraptor sp. along with (?)Conchoraptor specimen ZPAL MgD-I/95 and another specimen. Maryanska and Osmólska (1997) later described a fragmentary skull from the same locality, referring to it as ZPAL MgD-I/96. The identical locality and preserved material, along with the closeness in specimen number to ZPAL MgD-I/95 lead me to believe this is the same specimen Osmólska mentioned. It is described as crested, so may belong to Rinchenia or Nemegtomaia, both also from the Nemegt Formation.
980807 GT Coy Oviraptr (a field number for a joint HMNS-IGM collection) was found in August 7 1998 at Gurilin Tsav, and was said along with 980807 GT SZK Oviraptr (now IGM 102/12) to "belong to a single species, which is similar to the partial skeleton of oviraptorosaurians found at Bugin Tsav in 1994 by our party" (Suzuki and Watabe, 2000). It was later photographed as figure 18 of Tsogtbaatar and Chinzorig (2010) once prepared, which shows differs from Funston's Guriliin Tsav oviraptorid (including IGM 102/12) in having a shallower postacetabular process with greater invagination of the ischial peduncle, so is a different taxon.
Currie (2001) listed "Oviraptorid, partial skeleton" as excavated September 10-24 2000 from Khermeen Tsav, Nemegt Formation, so probably the Upper White Beds.
Weishampel et al. (2008) described a nest of four embryos which they stated resembled Nemegtomaia slightly more.
Currie et al.'s (2008) material is from an Avimimus bonebed, who noted less than ten percent was from other genera including oviraptorids.
References- Osmólska, 1976. New light on skull anatomy and systematic position of Oviraptor. Nature. 262, 683-684.
Maryanska and Osmólska, 1997. The quadrate of oviraptorid dinosaurs. Acta Palaeontologia Polonica. 42, 377-387.
Suzuki and Watabe, 2000. Report on the Japan-Mongolia Joint Paleontological Expedition to the Gobi desert, 1998. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 83-98.
Weishampel, Fastovsky, Watabe, Barsbold and Tsogtbaatar, 2000. New embryonic and hatchling dinosaur remains from the Late Cretaceous of Mongolia. Journal of Vertebrate Paleontology. 20(3), 78A.
Currie, 2001. Nomadic Expeditions, Inc. report on fieldwork in Mongolia, September 2000. Alberta Palaeontological Society, Fifth Annual Symposium. 12-16.
Badamkhatan, 2008. Dinosaurs from the Late Cretaceous Mongolian locality of Khaichin I. Journal of Vertebrate Paleontology. 28(3), 47A.
Currie, Longrich, Ryan, Eberth and Demchig, 2008. A bonebed of Avimimus sp. (Dinosauria: Theropoda) from the Late Cretaceous Nemegt Formation, Gobi desert: Insights into social behavior and development in a maniraptoran theropod. Journal of Vertebrate Paleontology. 28(3), 67A.
Weishampel, Fastovsky, Watabe, Varricchio, Jackson, Tsogtbaatar and Barsbold, 2008. New oviraptorid embryos from Bugin-Tsav, Nemegt Formation (Upper Cretaceous), Mongolia, with insights into their habitat and growth. Journal of Vertebrate Paleontology. 28(4), 1110-1119.
Tsogtbaatar and Chinzorig, 2010. Fossil specimens prepared in Mongolian Paleontological Center: 2002-2008. Hayashibara Museum of Natural Sciences Research Bulletin. 3, 155-166.

unnamed Oviraptoridae (Erben, 1995)
Late Cretaceous
Hugang Formation, Henan, China
Material- (STIPB coll.) eggshells
Comments- Wiemann et al. (2015) report several eggshells they refer to Macroolithus yaotuensis and Heyuannia, but the latter is based on the flawed reasoning of Cheng et al.. The latter assigned shells to Heyuannia "or an oviraptorosaurian of similar kind" based on its subarctometatarsaly. Yet this is true in most other oviraptorids as well except for "Ingenia".
References- Erben, 1995. The Cretaceous/Tertiary boundary in the Nanxiong-Basin (continental facies, SE-China). Stuttgart: Franz Steiner Verlag. 245 pp.
Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, Müller and Sander, 2015. The blue-green eggs of dinosaurs: How fossil metabolites provide insights into the evolution of bird reproduction. PeerJ PrePrints. DOI: 10.7287/peerj.preprints.1080v1
Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, Müller and Sander, 2017. Dinosaur origin of egg color: Oviraptors laid blue-green eggs. PeerJ. 5:e3706.

unnamed Oviraptoridae (Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, Müller and Sander, 2015)
Late Cretaceous
Nanxiong Group, Nanxiong Basin, Guangdong, China
Material- eggshells
Comments- Wiemann et al. (2015) report several eggshells they refer to Macroolithus yaotuensis and Heyuannia, but the latter is based on the flawed reasoning of Cheng et al. (2008).
References- Cheng, Ji, Wu and Shan, 2008. Oviraptorosaurian eggs (Dinosauria) with embryonic skeletons discovered for the first time in China. Acta Geologica Sinica. 82(6), 1089-1094.
Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, Müller and Sander, 2015. The blue-green eggs of dinosaurs: How fossil metabolites provide insights into the evolution of bird reproduction. PeerJ PrePrints. DOI: 10.7287/peerj.preprints.1080v1
Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, Müller and Sander, 2017. Dinosaur origin of egg color: Oviraptors laid blue-green eggs. PeerJ. 5:e3706.

unnamed Oviraptoridae (Sato, Cheng, Wu, Zelenitsky and Hsiao, 2005)
Cenomanian-Campanian, Late Cretaceous
Dafeng or Zhutian Formation, Nanxiong Group, Hongcheng Basin, Jiangxi, China
Material- (NMNS-CYN-2004-DINO-05/I) egg (Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, Müller and Sander, 2015)
Late Cretaceous
Nanxiong Group, Nankang District, Ganzhou, Jiangxi, China
(IVPP V20182) (embryo) partial skull (46.8 mm), partial mandible (38.7 mm), penultimate and last sacral vertebrae, sacral ribs, first to fourth caudal vertebrae, fifth caudal neural arch, proximal metatarsal II, proximal metatarsal III, proximal metatarsal IV, egg (198.3x88 mm) (Wang, Zhang, Sullivan and Xu, 2016)
(IVPP V20183) (embryo) incomplete nasal, lacrimal, frontal, anterior dentaries, three posterior sacral centra, mid sacral rib, first to sixth caudal centra, incomplete ilium (45.5 mm), incomplete pubes, partial ischium, femur (49.2 mm), tibia (56.2 mm), incomplete metatarsal II, phalanges II-?, incomplete metatarsal III, phalanges III-?, egg (163.5x74.8 mm) (Wang, Zhang, Sullivan and Xu, 2016)
(IVPP V20184) (embryo) partial parietal, incomplete squamosal, quadratojugal, quadrate, cervical centrum, two cervical neural arches, few dorsal centra, few fragmentary dorsal ribs, furcula, (?)femur, egg (179.5x92.1 mm) (Wang, Zhang, Sullivan and Xu, 2016)
Late Cretaceous
Nanxiong Group, Hongcheng Basin, Jiangxi, China
(Chimei Museum 41) (embryo) centra, distal femora, tibia, phalanges, unguals, fragments, partial egg (169x83.8 mm) (Cheng, Ji, Wu and Shan, 2008)
(NMNS-0015726-F02-embryo-01) (embryo) centra, neural arches, partial femora, tibiae (one incomplete; one partial), incomplete metatarsal II (~29.6 mm), metatarsal III (~32 mm), phalanx III-1, metatarsal IV (22.5 mm), phalanx IV-1, fragments, incomplete egg (173.5x76.3 mm) (Cheng, Ji, Wu and Shan, 2008)
(NMNS-VPDINO-2002-0901) six sacral vertebrae, first caudal vertebra, second caudal vertebra, incomplete ilia, proximal pubes, ischia, proximal femora, incomplete tibia, incomplete fibula, tarsus, two partial metatarsals, two eggs (175 mm) (Sato, Cheng, Wu, Zelenitsky and Hsiao, 2005)
Late Cretaceous
Nanxiong Group, Jiangxi, China
gastralia, posterior sacrum, over twenty-six caudal vertebrae, pelvis including ischia, hindlimb including femur, tibia and metatarsals, two eggs (193 mm) (He, Varricchio, Jackson, Jin and Pust, 2012)
Comments- The eggs of NMNS-VPDINO-2002-0901 are preserved inside the pelvic cavity and resemble Macroolithus yaotunensis and M. rugustus (Sato et al., 2005).
Cheng et al. (2008) found Chimei Museum 41 and NMNS-0015726-F02-embryo-01 were assignable to Macroolithus yaotunensis. They assigned these to Heyuannia "or an oviraptorosaurian of similar kind" based on its subarctometatarsaly. Yet this is true in most other oviraptorids as well except for H. yanshini, Gigantoraptor and "Tongtianlong", although not scorable in any genus except the latter from the Nanxiong formation.
He et al. (2012) remark on an additional specimen with two internal eggs, assignable only to Elongatoolithidae.
Wang et al. (2013) note embryos from the same formation inside of Macroolithus yaotunensis eggs. These are detailed in Wang et al. (2016), which only identifies the embryos as Oviraptoridae.
Wiemann et al. (2015) report several eggshells they refer to Macroolithus yaotuensis and Heyuannia, but the latter is based on the flawed reasoning of Cheng et al.. These are said to be from the "Nanxiong Formation of the Hongcheng area near Ganzhou City", which probably refers to the Hongcheng Basin. In Supplementary Table 2 of the published 2018 version, they are said to be from the Yuanpu Formation, which has since become the Dafeng and possibly the Zhutian Formation depending on whose stratigraphy the authors were using (Li et al. 2007 or Zhang et al 2013).
These remains may belong to the contemporaneous Banji, "Huanansaurus", "Corythoraptor", Jiangxisaurus, "Tongtianlong", Ganzhousaurus and/or Nankangia.
References- Sato, Cheng, Wu, Zelenitsky and Hsiao, 2005. A pair of shelled eggs inside a female dinosaur. Science. 308, 375.
Cheng, Ji, Wu and Shan, 2008. Oviraptorosaurian eggs (Dinosauria) with embryonic skeletons discovered for the first time in China. Acta Geologica Sinica. 82(6), 1089-1094.
He, Varricchio, Jackson, Jin and Pust, 2012. An oviraptorid adult-egg association and the origin of avialan reproductive strategies. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 108.
Wang, Zhang and Xu, 2013. New oviraptorid (Theropoda, Oviraptorosauria) embryos from the Upper Cretaceous of Southern China. Journal of Vertebrate Paleontology. Program and Abstracts 2013, 234.
Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, Müller and Sander, 2015. The blue-green eggs of dinosaurs: How fossil metabolites provide insights into the evolution of bird reproduction. PeerJ PrePrints. DOI: 10.7287/peerj.preprints.1080v1
Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, Müller and Sander, 2017. Dinosaur origin of egg color: Oviraptors laid blue-green eggs. PeerJ. 5:e3706.
Wang, Zhang, Sullivan and Xu, 2016. Elongatoolithid eggs containing oviraptorid (Theropoda, Oviraptorosauria) embryos from the Upper Cretaceous of southern China. BMC Evolutionary Biology. 16:67.

Beibeilong Pu, Zelenitsky, Lu, Currie, Carpenter, Xu, Koppelhus, Jia, Xiao, Chuang, Li, Kundrat and Shen, 2017
B. sinensis Pu, Zelenitsky, Lu, Currie, Carpenter, Xu, Koppelhus, Jia, Xiao, Chuang, Li, Kundrat and Shen, 2017
Coniacian, Late Cretaceous
Gaogou Formation, Henan, China
Holotype- (HGM 41HIII1219; formerly Children's Museum of Indianapolis coll.; Baby Louie) (embryo) incomplete skull, mandible (~65 mm), anterior cervical neural arch, anterior cervicodorsal centrum, last dorsal vertebra, six sacral vertebrae, few vertebrae, rib fragments, furcula, scapulae (one fragmentary), humeral shafts, partial radius, ulna (29 mm), metacarpal II? (15.2 mm), ilium (66 mm), pubes (62.4 mm), ischium, femora (~75 mm), proximal tibia, proximal fibula, phalanx II-1, phalanx III-2 (11.9 mm), phalanges IV-2 (6.1 mm), phalanx IV-3 (5.7 mm), phalanx IV-4, six-eight eggs (450, 400, 435 mm) one of which includes a limb element, nest (Currie, 1996)
....(LACM 7477/149736b) eggshells
Referred- (HX 9301) egg (Li, Yin and Liu, 1995)
(HX 9302) egg (Li, Yin and Liu, 1995)
(HX 9303) egg (Li, Yin and Liu, 1995)
(HXW 9301) egg (Li, Yin and Liu, 1995)
(HWX 9302) egg (Li, Yin and Liu, 1995)
(HWX 9303) egg (Li, Yin and Liu, 1995)
(LACM F.A.3818,2001-1) egg (Grellet-Tinner, 2005)
?(Nanyang Museum coll.) thirteen eggs, nest (Currie, 1996)
eggs (Wang and Zhou, 1995)
? eggs (Fang et al., 1998)
? eggs (Liang et al., 2009)
Diagnosis- (after Pu et al., 2017) antorbital fossa demarcated by sharply defined alveolar and dorsoposterior trending ridges; subantorbital portion of maxilla inset medially; posterodorsal margin of lacrimal overlapped by frontal; pronounced retroarticular process with distinct concave posterior facet (roughly as tall at the base as it is wide); preacetabular process of ilium longer than postacetabular process; posterior end of postacetabular process truncated or broadly rounded; accessory trochanter of femur weakly developed.
Comments- There is ambiguity in the formation the holotype was recovered from. Grellet-Tinner (2005) listed it as being from the Maastrichtian Zoumagang Formation, along with the Nanyang Museum nest and LACM F.A.3818,2001-1. Simon (2014) listed Wang and Zhou's (1995) material as being from the Cenomanian-Turonian Gaogou Formation, Liang et al.'s (2009) as being from either formation, and Fang et al.'s (1998) as being from the Zoumagang Formation, while Li et al.'s (1995) eggs were not specified as belonging to a particular formation. Pu et al. (2017) tracked the holotype locality to the Gaogou Formation, but whether Li et al.'s, Fang et al.'s, Liang et al.'s and/or Grellet-Tinner's materials are also actually from this formation is unclear. Pan et al. (2016) concur and narrow the Gaogou Formation's age to Coniacian.
The embryo nicknamed Baby Louie was discovered between December 1992 and early 1993 and donated to the Children's Museum of Indianapolis, though it was returned to China in 2013. Currie (1996) initially reported the specimen as a therizinosaur, since being reidentified as an oviraptorid (Norell and Clark, pers. comm. to Grellet-Tinner, 2005). Grellet-Tinner (2005) described the eggs in detail (published as Grellet-Tinner et al., 2006), though Baby Louie itself was not named or described in detail until Pu et al. (2017). The partial radius, ulna several vertebral centra and some rib fragments "are associated with another egg (#4) in the specimen, which may belong to the skeleton or to another perinate" (Pu et al., 2017).
Identical eggs from the same formation were described as the ootaxon Macroelongatoolithus xixiaensis (Li et al., 1995), senior synonym of Longiteresoolithus xixiaensis (Wang and Zhou, 1995) from the same formation. The former species has since been synonymized with M. carlylensis by Zelenitsky et al. (2000) and Simon (2014).
References- Li, Yin and Liu, 1995. The discovery of a new genus of dinosaur egg from Xixia, Henan, China. Journal of Wuhan Institute of Chemical Technology. 17, 38-41.
Wang and Zhou, 1995. The discovery of new typical dinosaur egg fossils in Xixia basin, Henan Province. Henan Geology. 13, 262-267.
Currie, 1996. The great dinosaur egg hunt. National Geographic. 189(May 1996), 96-111.
Fang, Lu, Cheng, Zou, Pang, Wang, Chen, Yin, Wang, Liu, Xie and Jin, 1998. On the Cretaceous fossil eggs of Xixia County, Henan Province. Geological Publishing House Beijing. 125 pp.
Zelenitsky, Carpenter and Currie, 2000. First record of elongatoolithid theropod eggshell from North America: The Asian oogenus Macroelongatoolithus from the Lower Cretaceous of Utah. Journal of Vertebrate Paleontology. 20(1), 130-138.
Grellet-Tinner, 2005. A phylogenetic analysis of oological characters: A case study of saurischian dinosaur relationships and avian evolution. PhD thesis, University of Southern California. 221 pp.
Grellet-Tinner, Chiappe, Norell and Bottjer, 2006. Dinosaur eggs and nesting behaviors: A paleobiological investigation. Palaegeography, Palaeoclimatology, Palaeoecology. 232, 294-321.
Liang, Wen, Yang, Zhou and Wu, 2009. Dinosaur eggs and dinosaur egg-bearing deposits (Upper Cretaceous) of Henan Province, Chian: Occurrences, palaeoenvirionments, taphonomy, and preservation. Progress in Natural Science. 19, 1587-1601.
Simon, 2014. Giant dinosaur (theropod) eggs of the oogenus Macroelongatoolithus (Elongatoolithidae) from southeastern Idaho: Taxonomic, paleobiogeographic and reproductive implications. Masters thesis, Montana State University. 110 pp.
Zelenitsky, Currie, Carpenter and Lu, 2015. Baby Louie: A theropod perinate from the Cretaceous of China reveals affinity of the largest known dinosaur eggs. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 244-245.
Pan, Xu, Pu, Jia, Lu, Zhou, Chang, Zhang and Yang, 2016. The geological age and paleoenvironment of Baby Louie bearing strata. Geological Bulletin of China. 35(12), 1961-1966.
Pu, Zelenitsky, Lu, Currie, Carpenter, Xu, Koppelhus, Jia, Xiao, Chuang, Li, Kundrat and Shen, 2017. Perinate and eggs of a giant caenagnathid dinosaur from the Late Cretaceous of central China. Nature Communications. 8:14952.

undescribed oviraptorid (Jensen, 1970)
Cenomanian, Late Cretaceous
Mussentuchit Member of Cedar Mountain Formation, Utah, US
Material- (BYU-E 200; holotype of Oolithes carlylensis) eggshell fragment (Jensen, 1970)
(BYU-E 201) eggshell fragment (Jensen, 1970)
(BYU VP13699; paratype of Oolithes carlylensis) eggshell fragment (Jensen, 1970)
(BYU VP13700; paratype of Oolithes carlylensis) eggshell fragment (Jensen, 1970)
(BYU VP13701; paratype of Oolithes carlylensis) eggshell fragment (Jensen, 1970)
(BYU VP13702; paratype of Oolithes carlylensis) eggshell fragment (Jensen, 1970)
(BYU VP13703) eggshell fragment (Jensen, 1970)
(BYU VP13704) eggshell fragment (Jensen, 1970)
(BYU VP13705) eggshell fragment (Jensen, 1970)
(BYU VP13706) eggshell fragment (Jensen, 1970)
(BYU coll.) >225 eggshell fragments (Jensen, 1970)
(TMP 1998.107.0004) eggshell fragment (Zelenitsky, Carpenter and Currie, 2000)
(TMP 1998.107.0007) eggshell fragment (Zelenitsky, Carpenter and Currie, 2000)
(TMP 1998.107.0010) eggshell fragment (Zelenitsky, Carpenter and Currie, 2000)
(TMP 1998.107.0015) eggshell fragment (Zelenitsky, Carpenter and Currie, 2000)
(TMP 1998.107.0018) eggshell fragment (Zelenitsky, Carpenter and Currie, 2000)
(TMP 1998.107.0022) eggshell fragment (Zelenitsky, Carpenter and Currie, 2000)
(TMP 1998.107 coll.) >1500 eggshell fragments (Zelenitsky, Carpenter and Currie, 2000)
(UCM 82008) eggshell fragment (Jensen, 1970)
partial skeleton including mid caudal vertebra and pygostyle (Makovicky, Shinya and Zanno, 2014)
Comments- The adult skeleton is said to be very large, "second only to ... Gigantoraptor in size." The mid caudal is "highly pneumatic" and the last four caudals "form a pygostyle-like structure." As the type material of Macroelongatoolithus carlylensis (Jensen, 1970) was found close by, it is hypothesized to belong to the species. This was originally described as Oolithes carlylensis by Jensen, then referred to Macroolithus by Zhao (1975), and named as the new genus Boletuoolithus by Bray (1998), before being referred to Macroelongatoolithus by Zelenitsky et al. (2000). M. carlylensis was emended to M. carlylei by Zelenitsky et al. (2000), but while the ICZN states species should originally be named this way (Article 31.1.3), it is not a spelling that must be corrected (Article 32.5).
References- Jensen, 1970. Fossil eggs in the Lower Cretaceous of Utah. Brigham Young University Research Studies. Geology Series. Geology Studies. 17, 51-65.
Zhao, 1975. The microstructure of the dinosaurian eggshells of Nanxiong basin, Guangdong province. Vertebrata PalAsiatica. 13, 105-117.
Bray, 1998. Dinosaur eggshell Boletuoolithus carlylensis, oogenus nov. from the Lower Cretaceous Cedar Mountain Formation of Utah. New Mexico Museum of Natural History and Science Bulletin. 14, 221-224.
Zelenitsky, Carpenter and Currie, 2000. First record of elongatoolithid theropod eggshell from North America: The Asian oogenus Macroelongatoolithus from the Lower Cretaceous of Utah. Journal of Vertebrate Paleontology. 20(1), 130-138.
Makovicky, Shinya and Zanno, 2014. New additions to the diversity of the Mussentuchit Member, Cedar Mountain Formation dinosaur fauna. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 175.

Gigantoraptor Xu, Tan, Wang, Zhao and Tan, 2007
G. erlianensis Xu, Tan, Wang, Zhao and Tan, 2007
Middle-Late Campanian, Late Cretaceous
Sanhangobi, Iren Dabasu Formation, Inner Mongolia, China
Holotype- (LH V0011) (8 m; ~2.1 tons; 11 year old adult) mandibles (~438 mm), posterior cervical neural arch, eight partial dorsal vertebrae, several dorsal ribs, gastralia, sacrum, twenty-seven caudal vertebrae, fourteen chevrons, incomplete scapula, furcula, sternum, humerus (735 mm), radius, ulna, scapholunare, semilunate carpal, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, proximal phalanx II-2, manual ungual II, distal metacarpal III, phalanx III-1, phalanx III-2, manual ungual III, partial ilium, pubes, femur (1.1 m), tibia (1.18 m), fibula, astragalus, calcaneum, metatarsus (583 mm), pedal phalanges
Diagnosis- (after Xu et al., 2007) mandible less than 45% of femoral length; fossa on the lateral surface of the dentary close to the anterior end; fossa bounded dorsally by a lateral flange anterodorsal to the external mandibular fenestra; long posteroventral process of the dentary extending to the level of the glenoid; small, posteriorly tapered retroarticular process much deeper than wide; opisthocoelous proximal caudal vertebrae; procoelous distal caudal vertebrae; pleurocoels present on most caudal vertebrae; pair of vertically arranged pneumatic openings present on the lateral surface of proximal caudal centra; large pneumatic opening present on the ventral surface of proximal and middle caudal centra; proximal caudal vertebrae with tall neural spines (about three times as tall as wide); proximal caudal vertebrae with robust and rod-like transverse processes located posteriorly; posteroventral margin of proximal caudal centra extending considerably ventrally; well-developed laminal system on the proximal caudal vertebrae (prespinal, postspinal, spinopostzygapophyseal, anterior centrodiapophyseal, posterior centrodiapophyseal, and prezygodiapophyseal laminae present); middle caudal vertebrae with vertical prezygapophyseal articular facets located proximal to the distal extremity of the process; prominent convexity ventral to the acromion process on the lateral surface of the scapula; laterally bowed humerus; humerus with prominent, spherical head; humerus with strongly medially curved deltopectoral crest; centrally constricted thick ridge running along the posterior margin of the proximal half of the humerus; ulna with a subcircular, concave proximal articular surface; radius with a subspherical distal end; metacarpal I with a slightly convex medial margin of the proximal end; medial condyle of metacarpal I three times as high as wide; medial condyle of metacarpal I extending much more distally than the lateral condyle; metacarpal II with prominent dorsolateral process on the proximal end; metacarpal II with longitudinal groove on the ventral margin of the proximal third of the shaft; manual unguals with a triangular set of lateral grooves; laterally compressed pubis; femur with straight shaft; constricted femoral neck; posteromedially oriented, spherical femoral head; anteroposteriorly wide trochanteric crest which is very robust and higher anteriorly than posteriorly; distinct narrow groove medial to the trochanteric crest extending down the posterior margin of the femoral shaft; patellar groove present on the anterior surface of the distal femur; small calcaneum obscured from anterior view by the wide astragalar main body; proximal projection on the lateral margin of distal tarsal IV; metatarsal III with ginglymoid distal end; pedal unguals with two lateral grooves; constricted proximal articular surface of pedal unguals.
Comments- Gigantoraptorwas discovered in April 2005 (Pickrell, 2014) and described by Xu et al. (2007). Additional elements were figured by Xing et al. (2012). This was found to be a basal oviraptorid by Xu et al. using Maryanska's oviraptorosaur matrix and a basal caenagnathid in a more recent version (Funston and Currie, 2016), a caenagnathid in Cau et al. (2017), and a basal oviraptorid in Hartman et al. (2019).
References- Xu, Tan, Wang, Zhao and Tan, 2007. A gigantic bird-like dinosaur from the Late Cretaceous of China. Nature. 844-847.
Xing, He, Li and Xi, 2012. A review on the study of the stratigraphy, sedimentology, and paleontology of the Iren Dabasu Formation, Inner Mongolia. In Dong (ed.). Proceedings of the Thirteenth Annual Meeting of the Chinese Society of Vertebrate Paleontology. China Ocean Press. 1-44.
Pickrell, 2014. Flying Dinosaurs: How fearsome reptiles became birds. NewSouth Publishing. 256 pp.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature. 552, 395-399. DOI: 10.1038/nature24679
Ma, Wang, Pittman, Tan, Tan, Guo and Xu, 2017. Functional anatomy of a giant toothless mandible from a bird-like dinosaur: Gigantoraptor and the evolution of the oviraptorosaurian jaw. Scientific Reports. 7: 16247. DOI:10.1038/s41598-017-15709-7
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

Nomingia Barsbold, Osmólska, Watabe, Currie and Tsogtbaatar, 2000
N. gobiensis Barsbold, Osmólska, Watabe, Currie and Tsogtbaatar, 2000
= "Nomingis brevicauda" Brush, DML 1999
= Nomingia "brevicaudia" Skrepnick, DML 2000
Early Maastrichtian, Late Cretaceous
Bugin Tsav, Nemegt Formation, Mongolia
Holotype- (IGM 100/119; listed as GIN 940824 by Barsbold et al. 2000; field number 940824 BgT MAT) (1.8 m; ~44 kg) last three cervical vertebrae, first-tenth dorsal vertebrae, ten fragmentary dorsal ribs, several gastralia, five sacral vertebrae, nineteen caudal vertebrae (460 mm; first caudal 28.3 mm), pygostyle (56 mm), fifteen chevrons, ilia (252 mm), pubes (243 mm), ischia (145 mm), femur (285 mm), tibiae (355 mm), fibulae, astragalus, calcaneum
Early Maastrichtian, Late Cretaceous
Nemegt, Nemegt Formation, Mongolia
Referred- (IGM 102/5; field number PJC.2001.5) pygostyle (Currie, 2002)
Diagnosis- (after Barsbold et al., 2000) twenty-four caudal vertebrae; last five caudal vertebrae fused; chevrons wide and dorsoventrally elongate on most caudals; preacetabular process ~25% longer than postacetabular process; dorsal ilial margin convex over preacetabular process and straight over postacetabular process; postacetabular process equally deep over entire length; pelvis weakly propubic, with pubic axis ~20 degrees to vertical; ilial peduncle of pubis twice as long as ilial pedunle of ischium.
Comments- The holotype was discovered in August 24 1994, prepared in 1997 (Tsogtbaatar, 2004), illustrated in Sloan (1999) and described briefly by Barsbold et al. (2000) before its official publication. Watabe and Suzuki (2000) provide a photo of the specimen in situ as "Postcranial skeleton of Oviraptorosauria from Bugin Tsav (Nemegt Formation)" and list it as "Oviraptorosauria whole skeleton without skull" with its field number. The taxon was originally called "Nomingia brevicaudia" (initially misspelled online as "Nomingis brevicauda" by the Dinosauricon e.g Brush, DML 1999), but was changed shortly before publication (Skrepnick, DML 2000). The latter species name has never been published however. Funston et al. (2018) proposed it may be synonymous with the contemporaneous Elmisaurus rarus, but combining the OTUs takes 7 more steps in Hartman et al.'s (2019) analysis.
Barsbold et al. (2000) did not place Nomingia in a family, but stated the long preacetabular process, straight pubis and strong posterior curve to the ischium might suggest caenagnathid affinities. Maryanska et al. (2002) first included it in a phylogenetic analysis and found Nomingia to be placed in the Caenagnathidae, but future analyses such as a more recent Maryanska derivative Funston and Currie (2016), Cau et al. (2017) and Hartman et al. (2019) have recovered it as a basal oviraptorid.
The holotype was a subadult based on the unfused sutures between presacral neural arches and centra. Based on comparison to oviraptorids and adjusting for the shorter tail, it may have been about 1.8 meters long.
References- Brush, DML 1999. https://web.archive.org/web/20210603191140/http://dml.cmnh.org/1999Nov/msg00038.html
Sloan, 1999. Feathers for T. rex? National Geographic. 196(5), 98-107.
Barsbold, Currie, Myhrvold, Osmólska, Tsogtbaatar and Watabe, 2000. A pygostyle from a non-avian theropod. Nature. 6766, 155
Barsbold, Osmólska, Watabe, Currie and Tsogtbaatar, 2000. A new oviraptorosaur (Dinosauria, Theropoda) from Mongolia: The first dinosaur with a pygostyle. Acta Paleontologica Polonica. 45(2), 97-106.
Skrepnick, DML 2000. https://web.archive.org/web/20210605025348/http://dml.cmnh.org/2000Oct/msg00074.html
Watabe and Suzuki, 2000. Report on the Japan-Mongolia Joint Paleontological Expedition to the Gobi desert, 1994. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 30-44.
Currie, 2002. Report on fieldwork in Mongolia, September 2001. Alberta Palaeontological Society, sixth annual symposium. 8-12.
Maryanska, Osmólska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Tsogtbaatar, 2004. Fossil specimens prepared in Mongolian Paleontological Center 1993-2001. Hayashibara Museum of Natural Sciences Research Bulletin. 2, 123-128.
Persons, Currie and Norell, 2011. Shake your feathers: The flamboyant, athletic, and possibly flirtatious caudal morphology of oviraptorosaurs. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 174.
Persons, Currie and Norell, 2014 (online 2013). Oviraptorosaur tail forms and functions. Acta Palaeontologica Polonica. 59(3), 553-567.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature. 552, 395-399. DOI: 10.1038/nature24679
Funston, Mendonca, Currie and Barsbold, 2018 (online 2017). Oviraptorosaur anatomy, diversity and ecology in the Nemegt Basin. Palaeogeography, Palaeoclimatology, Palaeoecology. 494, 101-120.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

Banji Xu and Han, 2010
B. long Xu and Han, 2010
Late Cretaceous
Nanxiong Group, Hongcheng Basin, Jiangxi, China
Holotype- (IVPP V16896) (juvenile) skull (65 mm), mandibles (one partial, one incomplete)
Diagnosis- (after Xu and Han, 2010) premaxillonasal crest with stepped posterior end; premaxillonasal crest with two longitudinal grooves and numerous oblique striations on lateral surface; elongate external naris placed posteriorly; deep fossa on dorsal surface of palatal process of pterygoid; several longitudinal grooves on the posterodorsal dentary; several tubercles on the dorsal surangular shelf.
Comments- Banji was discovered by an amateur collector prior to September 2009. Xu and Han (2010) initially recovered Banji as a basal oviraptorid based on a Maryanska et al. analysis, while a more modern version (Funston and Currie, 2016) moves it closer to heyuannines than Citipati or Rinchenia, sister to Wulatelong. Hartman et al. (2019) recover it in the more basal position however.
Reference- Xu and Han, 2010. A new oviraptorid dinosaur (Theropoda: Oviraptorosauria) from the Upper Cretaceous of China. Vertebrata PalAsiatica. 48(1), 11-18.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

"Huanansaurus" Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015
"H. ganzhouensis" Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015
Late Cretaceous
Nanxiong Group, Ganzhou Railway Station, Zhanggong District, Ganzhou, Jiangxi, China
Material- (HGM 41HIII-0443) incomplete skull (206.1 mm), mandible (182.8 mm), atlas, axis, third cervical vertebra, fourth cervical vertebra, fifth cervical vertebra, incomplete sixth cervical vertebra, partial seventh cervical vertebra, gastralia, incomplete humerus (195 mm), fragmentary radius (190 mm), fragmentary ulna, fragmentary carpals, fragmentary metacarpal I (40 mm), phalanx I-1 (80 mm), incomplete manual unguals I (70 mm), incomplete metacarpal II (85 mm), phalanges II-1 (one partial; 55 mm), phalanges II-2 (65 mm), manual unguals II (65 mm), incomplete metacarpal III (84 mm), phalanx III-1 (32 mm), phalanx III-2 (30 mm), phalanges III-3 (44 mm), manual unguals III (52 mm), distal femur, proximal tibia, incomplete metatarsal II, phalanx II-1 (45 mm), phalanx II-2 (35 mm), pedal ungual II (55 mm), incomplete metatarsal III, phalanx III-1 (45 mm), phalanx III-2 (39 mm), phalanx III-3 (35 mm), pedal ungual III (50 mm), incomplete metatarsal IV, phalanx IV-1 (33 mm), phalanx IV-2 (30 mm), phalanx IV-3 (21 mm), phalanx IV-4 (20 mm), pedal ungual IV (50 mm)
Diagnosis- (after Lu et al., 2015) posterodorsal process of the premaxillae contact lacrimals (also in Citipati and Conchoraptor); distinct opening near posteroventral corner of distal end of posterodorsal premaxillary process; circular supratemporal fenestra much smaller than laterotemporal fenestra; nuchal crest pronounced; mandibular condyles of quadrate posterior to occipital condyle; pneumatized dentaries; anterodorsal tip of dentary projecting anterodorsally at an angle of 45 degrees or less relative to the ventral margin of symphysis; length of dentary symphysis between 20% and 25% of mandible length; dentary portion of dorsal external mandibular fenestra margin strongly concave; posteroventral dentary process twisted so that lateral surface faces somewhat ventrally; angular contributes extensively to border of external mandibular fenestra; metacarpal I long and slender, diameter 20% of length; proximodorsal lip on all manual unguals prominent.
Comments- Lu et al. (2015) state "Huanansaurus" "was unearthed from the current construction site of the Ganzhou Railway Station." It was described on July 2 2015 as a new taxon of oviraptorid. However, this paper has no mention of ZooBank and as of February 6 2020 "Huanansaurus" lacks an entry on the ZooBank website. Thus according to ICZN Article 8.5.3 (an electronic work must "be registered in the Official Register of Zoological Nomenclature (ZooBank) (see Article 78.2.4) and contain evidence in the work itself that such registration has occurred"), "Huanansaurus ganzhouensis" Lu et al., 2015 is a nomen nudum that will only be technically valid pending action on behalf of the authors or ICZN as its journal is not published physically.
Lu et al. (2014) announced this specimen in an abstract as a sister taxon to Citipati. It was fully described and named by Lu et al. (2015), and emerged in a trichotomy with Citipati Osmólskae and IGM 100/42 in their analysis derived from Maryanska et al.'s oviraptorosaur matrix. Cau et al. (2017) also recovered it sister to C. Osmólskae (but not IGM 100/42) while Hartman et al. (2019) find it more basal.
References- Lu, Kobayashi, Pu, Chang, Zhang, Shang and Liu, 2014. A new oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China and its paleogeographical implications. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 164.
Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015. A new oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China and its paleobiogeographical implications. Scientific Reports. 5, 11490.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature. 552, 395-399. DOI: 10.1038/nature24679
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

unnamed oviraptorid "NXMV" (Lu, 2004)
Early Maastrichtian, Late Cretaceous
Zhenshui Formation, Nanxiong Group, Xincheng area, Nanxiong City, Guangdong, China
Material- (E-1) partial skull, mandible (150 mm)
....(K2-1) two posterior cervical vertebrae (30 mm), nine dorsal vertebrae (20 mm), dorsal rib fragment
....(K2-12) incomplete pes
Comments- These specimens probably belong to the same individual, called NXMV by Lu (2004). While Lu states it "came from the top of the Pingling Member of Shanghu Formation (Zhang, Personnel communication, 2002) or the top of the Pingling Formation (Zhao, personnel communication, 2002)", the Pingling Formation was replaced by the Zhenshui Formation by Li et al. 2007, dated as early Maastrichtian by Xi et al. 2021.
Wang et al. (2013) believes it may belong to Ganzhousaurus, but Hartman et al. (2019) recover it as sister to "Corythoraptor" in Oviraptoridae, while Ganzhousaurus is a non-caenagnathoid. Adding additional taxa leaves it in the same area of the tree, but in an uncertain position relative to "Corythoraptor", Nankangia, "Lingyuanosaurus" and Yulong.
References- Lu, 2004. Oviraptorid dinosaurs from southern China. PhD thesis, Southern Methodist University. 249 pp.
Lu, 2005. Oviraptorid Dinosaurs from Southern China. Geological Publishing House, Beijing. 200 pp.
Wang, Sun, Sullivan and Xu, 2013. A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of southern China. Zootaxa. 3640(2), 242-257.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

"Corythoraptor" Lu, Li, Kundrat, Lee, Sun, Kobayashi, Shen, Teng and Liu, 2017
"C. jacobsi" Lu, Li, Kundrat, Lee, Sun, Kobayashi, Shen, Teng and Liu, 2017
Late Cretaceous
Nanxiong Group, Ganzhou Railway Station, Zhanggong District, Ganzhou, Jiangxi, China
Material- (JPM-2015-001) (>7 year old adult) incomplete skull (~174.4 mm), incomplete mandibles (~143.2 mm), hyoids, atlas, axis (52 mm without odontoid), third cervical vertebra (55.3 mm), fourth cervical vertebra (67.8 mm), fifth cervical vertebra (66.4 mm), sixth cervical vertebra (71.7 mm), seventh cervical vertebra (62.6 mm), eighth cervical vertebra (63.7 mm), ninth cervical vertebra (64.5 mm), tenth cervical vertebra (65.7 mm), eleventh cervical vertebra (71.4 mm), twelfth cervical vertebra (53.2 mm) (dorsal series ~400 mm) first through sixth dorsal vertebrae, several dorsal ribs, gastralia, sacrum (~210 mm), first through fifth caudal vertebrae, dorsal third chevron, distal scapula, sternal plate, humerus (212.9 mm), radii (205 mm), ulnae (210 mm), carpals, metacarpals I (39.4 mm), phalanges I-1 (69.5 mm), manual unguals I (~60.5 mm), metacarpals II (96.5 mm), phalanges II-1 (64.5 mm), phalanges II-2 (63.5 mm), manual ungual II (60.9 mm), metacarpals III (93.8 mm), phalanges III-1 (36.3 mm), phalanx III-2 (31.1 mm), phalanges III-3 (50.9 mm), manual unguals III (55.6 mm), partial ilia (300 mm), pubes (360 mm), ischia (220 mm), femora (330.5 mm), tibiae (one incomplete; 392 mm), proximal fibuila, partial astragalus, distal tarsal III, distal tarsal IV, metatarsal I, phalanx I-1 (37.2 mm), pedal ungual I (36.9 mm), metatarsals II (one incomplete; 148.3 mm), phalanges II-1 (49.2 mm), phalanges II-2 (30.4 mm), pedal unguals II (52.1 mm), metatarsals III (157 mm), phalanges III-1 (48 mm), phalanges III-2 (30.5 mm), phalanges III-3 (30.5 mm), pedal unguals III (54.2 mm), metatarsals IV (156 mm), phalanges IV-1 (33.2 mm), phalanges IV-2 (24.4 mm), phalanges IV-3 (22.9 mm), phalanges IV-4 (28.9 mm), pedal unguals IV (49.6 mm), metatarsal V (45 mm)
Diagnosis- (after Lu et al., 2017) ratio of length of tomial margin of premaxilla to premaxilla height (ventral to the external naris) 1.0-1.4; inclination of anteroventral margin of premaxilla relative to horizontally positioned jugal ventral margin posterodorsal; dorsal process of premaxilla bears two processes- a short posterodorsally extending process, forming the anterodorsal margin of the external naris, and a long process, forming most of the anterodorsal process of the premaxilla; distinct cassowary-like helmet on skull; external naris much longer than tall; long axis of external naris parallel to dorsal margin of antorbital fenestra; antorbital fossa bordered anteriorly by maxilla; infratemporal fenestra dorsoventrally elongate, narrow anteroposteriorly; straight anterodorsal margin of dentary in lateral view; deep fossa, sometimes with associated pneumatopore, on lateral surface of dentary; no pleurocoels on second through fourth cervical vertebrae; length of neck twice as long as dorsal vertebral column, and slightly longer than forelimb length; less pronounced deltopectoral crest, forming an arc rather than being quadrangular; manus / humerus + radius ratio between 0.50 and 0.65; manual ungual III less curved than other unguals; anterior trochanter completely fused with greater trochanter; distal ends of metatarsal II straight and metatarsal IV laterally deflected.
Comments- Discovered in 2013 (Geggel, 2017), this was later described by Lu et al. on July 27 2017 as a new taxon of oviraptorid. However, this paper has no mention of ZooBank and as of February 6 2020 "Corythoraptor" lacks an entry on the ZooBank website. Thus according to ICZN Article 8.5.3 (an electronic work must "be registered in the Official Register of Zoological Nomenclature (ZooBank) (see Article 78.2.4) and contain evidence in the work itself that such registration has occurred"), "Corythoraptor jacobsi" Lu et al., 2017 is a nomen nudum that will only be technically valid pending action on behalf of the authors or ICZN as its journal is not published physically..
Lu et al. (2017) recovered this as sister to "Huanansaurus", similar to where Hartman et al. (2019) found it.
References- Geggel, 2017. Newfound dino looks like the creepy love child of a turkey and an ostrich. Live Science. https://www.livescience.com/59958-newfound-dinosaur-has-cassowary-like-crest.html
Lu, Li, Kundrat, Lee, Sun, Kobayashi, Shen, Teng and Liu, 2017. High diversity of the Ganzhou oviraptorid fauna increased by a new "cassowary-like" crested species. Scientific Reports. 7: 6393. DOI: 10.1038/s41598-017-05016-6
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

Nankangia Lu, Yi, Zhong and Wei, 2013
N. jiangxiensis Lu, Yi, Zhong and Wei, 2013
Late Cretaceous
Nanxiong Group, Longling Town, Nankang District, Ganzhou, Jiangxi, China
Holotype- (GMNH F10003) anterior mandibles, five mid dorsal vertebrae (38, 37, 38, 39, 40 mm), partial penultimate sacral vertebra, last sacral vertebra (43 mm), three proximal caudal vertebrae (38, 44, 42 mm), proximal caudal neural arch, six mid caudal centra (41, 39, 38, 41, 38, 37 mm), incomplete mid caudal vertebra, two mid chevrons, scapulocoracoids (scapula 260 mm), furcular fragments, incomplete humerus (240 mm), ilia (one incomplete; 330 mm), pubes (one incomplete; 380 mm), ischia (205 mm), femora (350, 380 mm), tibia (400 mm), fibular fragment, astragalus, calcaneum
Diagnosis- (after Lu et al., 2013) rostral end of mandibular symphyseal region not downturned (also in caenagnathids, Incisivosaurus, Luoyanggia and Ganzhousaurus); two infradiapophyseal fossae on ventral surface near base of transverse process of dorsal vertebrae; pneumatic fossae on sacral vertebrae slit-like; neural spines of proximal caudal vertebrae wider transversely than anteroposteriorly, forming a large posterior fossa with rugose central area; large fossa on anterior surface (infraprezygapophyseal fossa) and another (infradiapophyseal fossa) on ventral surface of base of transverse process of the proximal caudals; femur longer than ilium (also in Yulong and Khaan); ratio of height to length of ilium 0.36; femoral neck extending dorsomedially at about an angle of 90 degrees to the shaft; femur and tibia approximately the same length.
Comments- The holotype was donated to the GMNH by a farmer in 2010. Lu et al. (2013) added Nankangia to a version of the Maryanska et al. analysis and found it to be the basalmost oviraptorid except Gigantoraptor, but incorrectly placed Oviraptoridae at a less inclusive node. A more recent version (Funston and Currie, 2016) finds Nankangia as the most basal oviraptorid, having moved Gigantoraptor to Caenagnathidae. Cau et al. (2017) recovers it in Caenagnathidae while Hartman et al. (2019) find it to be a basal oviraptorid.
References- Lu, Yi, Zhong and Wei, 2013. A new oviraptorosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China and its paleoecological implications. PLoS ONE. 8(11), e80557.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature. 552, 395-399. DOI: 10.1038/nature24679
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

"Lingyuanosaurus" Yao, Liao, Sullivan and Xu, 2019
"L. sihedangensis" Yao, Liao, Sullivan and Xu, 2019
Early Albian, Early Cretaceous
Sihedang, Jiufotang Formation, Liaoning, China
Material- (IVPP V23589) (~12 kg, juvenile) cervical centrum (~30 mm), posterior dorsal neural arch, several dorsal ribs, mid sacral centrum (~25 mm), proximal caudal vertebra (~15 mm), three partial vertebrae, proximal humerus, distal humerus, manual ungual II, manual ungual III, ilium (143 mm), proximal (?)ischium, femur (200 mm), incomplete tibia, partial astragalus, fragments
Diagnosis- (after Yao et al., 2019) posterior dorsal vertebrae with prominent paradiapophyseal lamina separating anterior and posterior infradiapophyseal fossae; ventral part of hyposphene expanding transversely to form intumescence which extends as far as postzygapophysis posteriorly and beyond postzygapophysis laterally; dorsal centroprezygapophyseal lamina extending anteriorly considerably beyond level of prezygapophysis and fusing with opposite lamina along midline; ilium with most of dorsal margin strongly convex but posterior part of dorsal margin distinctly concave; subtriangular cuppedicus fossa immediately dorsal to pubic peduncle on medial surface of ilium.
Comments- "Lingyuanosaurus" was discovered before July 2018 and described on March 22 2019 in Scientific Reports. However, this paper has no mention of ZooBank and as of February 5 2020 "Lingyuanosaurus" lacks an entry on the ZooBank website. Thus according to ICZN Article 8.5.3 (an electronic work must "be registered in the Official Register of Zoological Nomenclature (ZooBank) (see Article 78.2.4) and contain evidence in the work itself that such registration has occurred"), "Lingyuanosaurus sihedangensis" Yao et al., 2019 is a nomen nudum that will only be technically valid pending action on behalf of the authors or ICZN as its journal is not published physically.
Yao et al. (2019) analyzed "Lingyuanosaurus" in a version of Zanno's therizinosaur matrix and resolved it between Beipiaosauirus and Alxasaurus as a therizinosauroid. However, when Mortimer added the taxon to Hartman et al.'s (2019) analysis, it emerged as an oviraptorid. Of the supposedly therizinosaurian characters listed by Yao et al., Zanno's "dorsal vertebrae with a complex laminar structure" refers to mid-dorsals ("anterior dorsals" in Zanno's nomenclature) which are not preserved in "Lingyuanosaurus" and nor was that character (their 274) scored for the taxon. Oviraptorosaurs can have both "laterally flattened manual unguals with dorsally positioned collateral grooves" (e.g. Chirostenotes) and "a highly modified ilium with a deep preacetabular process, a reduced postacetabular process, a preacetabular process whose ventral margin is dorsally displaced relative to the acetabulum" (e.g. Rinchenia). The slender pubic peduncle is therizinosaur-like, however. It only takes one more step to move the taxon to Therizinosauria, and four more to move to Ornithomimosauria. Notably, Zanno's oviraptorosaur sample was limited due to Balanoff working on the group at the time.
References- Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
Mortimer, 2019. https://theropoddatabase.blogspot.com/2019/03/what-is-lingyuanosaurus.html
Yao, Liao, Sullivan and Xu, 2019. A new transitional therizinosaurian theropod from the Early Cretaceous Jehol Biota of China. Scientific Reports. 9:5026. DOI: 10.1038/s41598-019-41560-z

Yulong Lu, Currie, Xu, Zhang, Pu and Jia, 2013
Y. mini Lu, Currie, Xu, Zhang, Pu and Jia, 2013
Late Maastrichtian, Late Cretaceous
Qiupa Formation, Henan, China
Holotype- (HGM 41HIII-0107) (<1 year old young juvenile) skull (48.16 mm), mandibles (51.97 mm), atlas, axis, five cervical vertebrae, posterior dorsal vertebrae, dorsal ribs, about twenty proximal and mid caudal vertebrae, scapula (42 mm), clavicles, humeri (44.6 mm), radii (44.5 mm), ulnae (44.7 mm), distal phalanx I-1, manual ungual I, metacarpal II (22.8 mm), phalanx II-1, phalanx II-2, manual ungual II, metacarpal III, phalanx III-2, phalanx III-3, manual ungual III, incomplete ilium (55 mm), partial femur (72 mm), tibiae (88.8 mm), incomplete fibulae, astragali, metatarsal I, partial phalanx I-1, metatarsals II, metatarsals III (43.3 mm), metatarsals IV, pedal phalanges including III-3 (11.7 mm), pedal unguals
Paratypes- (HGM 41HIII-0301) (embryo) skeleton including radius (22.6 mm), ulna (21 mm), ilium (39.3 mm), femur (46 mm), 26 eggs (19 not purchased), nest
(HGM 41HIII-0108) (young juvenile) posterior skull, posterior mandibles
(HGM 41HIII-0109) (young juvenile) skull, mandibles (48.54 mm), partial postcranial skeleton including scapula (50.26 mm), humerus (35.9 mm)
(HGM 41HIII-0110) (young juvenile) incomplete skull, incomplete mandibles, few cervical centra
(HGM 41HIII-0111) (young juvenile) ilium
Diagnosis- (after Lu et al., 2013) posterodorsal corner of antorbital fenestra and anteroventral corner of external naris at the same level (also in Citipati); distinct opening in premaxilla anteroventral to the external naris; antorbital fossa partly bordered by premaxilla anterodorsally; subnarial process of premaxilla does not contact anterior process of lacrimal; parietal almost as long as frontal; in dorsal view, posterior margin forms a straight line between postzygapophyses in fourth and fifth cervical vertebrae; femur longer than ilium.
Comments- Kobayashi et al. (2008) first mentioned Qiupa oviraptorids, and the Yulong material was found before 9-20-2012. While the holotype and HGM 41HIII-0108 were found associated, the others were found in separate quarries up to 4 km apart. The diagnostic characters may merely be characteristic of juvenile oviraptorids, and Yulong's recovered poisition in Lu et al.'s (2013) Maryanksa et al.-based analysis (between Gigantoraptor and more derived oviraptorids) may be basal due to ontogenetically variable characters, as juvenile theropods commonly end up more basally than adults.
References- Kobayashi, Lu, Lee, Xu and Zhang, 2008. A new basal ornithomimid (Dinosauria: Theropoda) from the Late Cretaceous in Henan province of China. Journal of Vertebrate Paleontology. 28(3), 101A.
Lu, Currie, Xu, Zhang, Pu and Jia, 2013. Chicken-sized oviraptorid dinosaurs from central China and their ontogenetic implications. Naturwissenschaften. 100(2), 165-175.

Oviraptorinae Barsbold, 1976a sensu Barsbold, 1981
Definition- (Oviraptor philoceratops + Citipati Osmólskae) (Osmólska, Currie and Barsbold, 2004)
Comments- Barsbold created this taxon to separate Oviraptor (to which he referred specimens now distinguished as Conchoraptor and Citipati? sp.) from "Ingenia". Once he named Conchoraptor and Rinchenia (originally Oviraptor mongoliensis), these were placed in Oviraptorinae as well. Barsbold et al. (1990) retained this taxonomy, though their cladogram shows oviraptorines to be paraphyletic to "ingeniines", with Conchoraptor closer to "Ingenia" than to Oviraptor (still including Citipati? sp.). Similarly, Maryanska et al. (2002) recovered Rinchenia, Citipati and Conchoraptor as paraphyletic to "Ingenia". The definition of Osmólska et al. (2004) functions in their phylogeny, where Rinchenia and Citipati form a clade exclusive of "Ingenia", Conchoraptor and Khaan. Oviraptor was not included in the analysis, but was assumed to be part of this clade, perhaps due to its cranial crest. Heyuannia was assigned to Oviraptorinae in their taxon list, but not included in the analysis, nor was the rationale for the assignment discussed. It's apparent the original concept of Oviraptorinae is probably not monophyletic with respect to "Ingeniinae", and using Osmólska et al.'s definition leads to situations where useful similarity to their or Barsbold's concept of the taxon is lost. A more useful definition would be (Oviraptor philoceratops <- Heyuannia yanshini), or with the recent replacement of "Ingeniinae" with Heyuanninae (Oviraptor philoceratops <- Heyuannia huangi).
References- Barsbold, 1976a. O novum pozdnemelovom semeystve melkikh teropod (Oviraptoridae fem. n.) Mongolii. Doklady Akademii Nauk SSSR. 226(3), 685-688.
Barsbold, 1976b. A new Late Cretaceous family of small theropods (Oviraptoridae n. fam.) in Mongolia. Doklady Akademia Nauk SSSR. 226, 221-223.
Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological Expedition Transactions. 15, 28-39.
Barsbold, Maryanska and Osmólska, 1990. Oviraptorosauria. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria. University of California Press. 249-258.
Maryanska, Osmólska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Osmólska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel, Dodson and Osmólska, (eds.). The Dinosauria, Second Edition. University of California Press. 165-183.

Citipatiinae Funston, Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020
Comments- Funston et al. (2020) used a version of Maryanska et al.'s oviraptorosaur analysis to recover "Nankangia jiangxiensis, Oviraptor philoceratops, and Yulong mini as successive outgroups to two subfamilies of oviraptorids." "One of the two groups is comprised of forms with elongate manus with digits roughly equal in robustness and an elongate third digit" which "is hereby designated Citipatiinae because Citipati is the oldest valid genus named in the family [sic]." While not explicitly listed as fam. nov. as suggested by ICZN Recommendation 16A, the authors do later refer to "this new citipatiine clade", which is considered to cover ICZN Article 16.1 ("Every new name published after 1999, including new replacement names (nomina nova), must be explicitly indicated as intentionally new"). While no phylogenetic definition is given, the wording and symbols in their Figure 6 suggests "Citipati Osmólskae <- Heyuannia huangi" was the intention.
Reference- Funston, Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020. A new two-fingered dinosaur sheds light on the radiation of Oviraptorosauria. Royal Society Open Science. 7: 201184.

"Anomalipes" Yu, Wang, Chen, Sullivan, Wang, Wang and Xu, 2018
"A. zhaoi" Yu, Wang, Chen, Sullivan, Wang, Wang and Xu, 2018
Campanian, Late Cretaceous
Xingezhuang or Hongtuya Formation, Wangshi Group, Shandong, China
Material- (ZCDM V0020) incomplete femur (~300 mm), incomplete tibia (~360 mm), partial fibula, pedal ungual II (51.4 mm), metatarsal III (167 mm), phalanx IV-1 (37.4 mm)
Diagnosis- (after Yu et al., 2018) femoral head anteroposteriorly narrow and somewhat deflected posteriorly; accessory trochanter low; lateral ridge present on femur; weak fourth trochanter present; metatarsal III with triangular proximal articular surface; prominent anterior flange near proximal end of metatarsal III; metatarsal III medial condyle much narrower than lateral condyle; longitudinal groove on distal articular surface of metatarsal III; pedal ungual II with lateral collateral groove deeper and more dorsally located than medial groove.
Comments- This was described by Yu et al. on March 22 2018 as a new taxon of caenagnathid. However, this paper has no mention of ZooBank and as of February 6 2020 "Anomalipes" lacks an entry on the ZooBank website. Thus according to ICZN Article 8.5.3 (an electronic work must "be registered in the Official Register of Zoological Nomenclature (ZooBank) (see Article 78.2.4) and contain evidence in the work itself that such registration has occurred"), "Anomalipes zhaoi" Yu et al., 2018 is a nomen nudum that will only be technically valid pending action on behalf of the authors or ICZN as its journal is not published physically.
Yu et al. (2018) used a version of Maryanska et al.'s oviraptorosaur analyusis to place this sister to Gigantoraptor in basal Caenagnathidae. In Hartman et al. (2019) it is an oviraptorid sister to Citipati.
References- Yu, Wang, Chen, Sullivan, Wang, Wang and Xu, 2018. A new caenagnathid dinosaur from the Upper Cretaceous Wangshi Group of Shandong, China, with comments on size variation among oviraptorosaurs. Scientific Reports. 8:5030. DOI: 10.1038/s41598-018-23252-2
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

Citipati Clark, Norell and Barsbold, 2001
Diagnosis- (after Clark et al., 2002) accessory opening on the lateral surface of the ascending process of the premaxilla.
C. Osmólskae Clark, Norell and Barsbold, 2001
Late Campanian, Late Cretaceous
Ukhaa Tolgod, Djadochta Formation, Mongolia
Holotype- (IGM 100/978) (~105 kg) complete skeleton including skull (160 mm), sclerotic ossicles, mandibles (151 mm), hyoids, twelve cervical vertebrae, cervical ribs, seven dorsal vertebrae, two uncinate processes, thirty caudal vertebrae (first caudal 38.2 mm), pygostyle, chevrons, scapulocoracoid, furcula, sternal plates, humerus (230 mm), radius (189 mm), ulna (200 mm), femur (345 mm), tibia (397 mm), astragalus, metatarsal I (37 mm), tarsometatarsus (II 176 mm; III 192 mm; IV 188 mm)
Paratypes- (IGM 100/971) (embryo) partial skull (~40 mm), mandibles, cervical vertebra, three fragmentary dorsal vertebrae, several dorsal rib fragments, three sacral centra, few distal caudal vertebrae, scapulae, coracoids, furcula, humeri (20 mm), proximal radius, proximal ulna, partial ilia, femur (36 mm), tibia (50 mm), fibula, astragali, partial metatarsal II, partial metatarsal III, partial metatarsal IV, pedal phalanges, nest, eggshell fragments (Norell et al., 1994)
?(IGM 100/979; Big Mama) distal cervical rib, anterior dorsal vertebra, distal dorsal ribs 1-6, four uncinate processes, gastralia, incomplete furcula, sternal plates, four sternal ribs, partial scapulae, coracoids, humeri (215 mm; one proximal), radii (198 mm; one distal), ulnae (214 mm; one distal), proximal carpal, semilunate carpal, metacarpal I (~45 mm), phalanx I-1 (88.4 mm), manual ungual I (91.1 mm), metacarpal II (89.5 mm), phalanx II-1 (65.5 mm), phalanx II-2 (61.5 mm), manual ungual II (76 mm), metacarpal III (89.4 mm), phalanx III-1 (33.8 mm), phalanx III-2 (~31 mm), phalanx III-3 (~45 mm), manual ungual III, distal pubes, distal ischia, distal femur, incomplete tibia, fibulae (~415 mm), astragalus, calcaneum, metatarsal I, phalanx I-1, pedal ungual I, metatarsal II (~173 mm), phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III (~190 mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV (~177 mm), phalanx IV-1, phalanx IV-2 (24.9 mm), phalanx IV-3 (20.1 mm), phalanx IV-4 (19 mm), pedal ungual IV (48.4 mm), nest, fifteen eggs (Clark et al., 1999)
Referred- ?(IGM 100/1004; Big Auntie) (13 year old adult) third to thirteenth cervical vertebrae fused with cervical ribs, ten dorsal vertebrae, ten dorsal ribs, four uncinate processes, at least four sacral vertebrae, incomplete scapulacoracoids (scapula 292.8 mm), furcula, sternal fragment, three sternal ribs, humeri (230.1 mm), radii (212.4 mm), ulnae (211.5 mm), scapholunare, semilunate carpal, phalanx I-1 (92.9 mm), manual ungual I (84.1 mm), metacarpal II (103.5 mm), phalanx II-1 (68.3 mm), phalanx II-2 (75.6 mm), manual ungual II, metacarpal III, phalanx III-1 (44.5 mm), phalanx III-2 (48.2 mm), phalanx III-3 (53.7 mm), manual ungual III (57.8 mm), partial ilium (~253 mm), proximal pubis, partial ischia, femora (one partial; ~402.4 mm), tibiae (one incomplete), fibulae (one incomplete), astragali, metatarsal I (42.4 mm), phalanx I-1 (31.9 mm), pedal ungual I (35.7 mm), distal metatarsal II, phalanx II-1 (53.2 mm), phalanx II-2 (37 mm), distal metatarsal III, distal metatarsal IV, phalanx IV-1 (38.6 mm), phalanx IV-2 (35.7 mm), phalanx IV-3 (31.6 mm), phalanx IV-4 (29 mm), pedal ungual IV (49.7 mm), twelve eggs (~181x~66.8 mm) (Webster, 1996)
(IGM 100/1009) (juvenile) skull (Norton, DML 2000)
? Late Campanian, Late Cretaceous
? Ukhaa Tolgod, Djadochta Formation, Mongolia
?(IGM 100/1125) two eggs (Grellet-Tinner, 2005)
Diagnosis- (after Clark et al., 2002) anterodorsally sloping occiput and quadrate; parietal much longer along the midline than the frontal and reaching nearly to the level of the anterior end of the orbit; ascending process of jugal perpendicular to the horizontal ramus (rather than extending posterodorsally); external naris nearly circular; nasal process of premaxilla vertical rather than sloping posterodorsally; cervical vertebrae elongate (approximately twice as long as they are wide).
Comments- The first published specimen of this species was an embryo discovered in 1993 in a nest of eggs (Norell et al., 1994), later described in detail by Norell et al. (2001). Originally assigned to Oviraptoridae indet., it was tentatively referred to the then unnamed Citipati Osmólskae by Norell et al. (2001) and formally referred to that taxon by Clark et al. (2001). Two partial juvenile paravian skulls were found with the specimen, originally identified as Velociraptor (Norell et al., 1994) and later troodontids (Norell and Makovicky, 1999).
Norell et al. (1995) announced a brooding oviraptorid on a nest found in 1993, which was initially referred to Oviraptor based on manual resemblences. It was mentioned in the popular press as "Big Mama". The specimen was later described in detail (Clark et al., 1999) and said to be an oviraptorid most closely related to Oviraptor, before being officially referred to Citipati Osmólskae by Clark et al. (2001).
A second brooding Citipati specimen found in 1995 is known as "Big Auntie" (Clark pers. comm. to Auditore). It was mentioned by Webster (1996) and Clark et al. (1999). A photograph of this specimen is present in Codd (2004), incorrectly identified as IGM 100/42. Similarly, it is photographed and incorrectly identified as Oviraptor philoceratops holotype AMNH 6517 by Codd et al. (2007). Erickson et al. (2009) examined it histologically, Grellet-Tinner (2005) described the eggs in detail in his thesis, and the specimen was fully described by Norell et al. (2018). The eggs are most similar to Elongatoolithus frustrabilis.
The holotype of the genus, a virtually complete specimen found in 1994, was initially identified as Oviraptor philoceratops (Webster, 1996) until it was preliminarily described and officially named by Clark et al. (2001). The skull and mandibles were later described in detail by Clark et al. (2002), though the postcrania remains largely undescribed and unillustrated, with only Nesbitt et al. (2009) and Balanoff and Norell (2012) showing the furcula and describing some details. More recently, Persons et al. (2014) describe the tail and pygostyle, while Norell et al. (2018) illustrate the third to fifth cervical vertebrae and two uncinate processes. Balanoff et al. (2018) described the endocranium based on Balanoff's (2011) thesis.
Norton (DML, 2000) noted IGM 100/1009 was discovered in 1993 at Ukhaa Tolgod and was on display at the AMNH Fighting Dinosaurs exhibit. Bhullar et al. (2012) listed it as a Citipati embryo.
References- Gibbons, 1994. Dino embryo recasts parents' image. Science. 266, 731.
Norell, Clark, Dashzeveg, Barsbold, Chiappe, Davidson, McKenna and Novacek, 1994. A theropod dinosaur embryo, and the affinities of the Flaming Cliffs dinosaur eggs. Science 266, 779-782.
Clark, 1995. The egg thief exonerated. Natural History. 6/95, 56-56.
Norell, Clark, Chiappe, and Dashzeveg, 1995. A nesting dinosaur. Nature. 378, 774-776.
Monastersky, 1996. Nesting dinosaur discovered in Mongolia. Science News. 149, 7.
Webster, 1996. Dinosaurs of the Gobi. National Geographic. 190(1), 70-89.
Norell and Clark, 1997. Birds are dinosaurs. Sci. Spectrum. 8, 28-34.
Clark, Norell and Chiappe, 1998. A "brooding" oviraptorid from the Late Cretaceous of Mongolia and its avian characters. Journal of Vertebrate Paleontology. 18(3), 34A.
Clark, Norell and Chiappe, 1999. An oviraptorid skeleton from the Late Cretaceous of Ukhaa Tolgod, Mongolia, preserved in an avianlike brooding position over an oviraptorid nest. American Museum Novitates. 3265, 1-36.
Norell and Makovicky, 1999. Important features of the dromaeosaur skeleton II: information from newly collected specimens of Velociraptor mongoliensis. American Museum Novitates. 3282, 1-45
Norell, Clark and Chiappe, 2001. An embryonic oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of Mongolia. American Museum Novitates. 3315 1-17.
Clark, Norell and Barsbold, 2001. Two new oviraptorids (Theropoda: Oviraptorosauria) from the Late Cretaceous Djadokta Formation, Ukhaa Tolgod. Journal of Vertebrate Paleontology. 21(2), 209-213.
Clark, Norell and Rowe, 2002 online. Citipati Osmólskae, Digital Morphology. http://digimorph.org/specimens/Citipati_Osmólskae/
Clark, Norell and Rowe, 2002. Cranial anatomy of Citipati Osmólskae (Theropoda, Oviraptorosauria), and a reinterpretation of the holotype of Oviraptor philoceratops. American Museum Novitates. 3364, 1-24.
Codd, 2004. The uncinate processes in birds and their implications for the breathing mechanics of maniraptoran dinosaurs. Dissertation zur Erlangung des Doktotgrades der Mathematisch-Naturwissenschaftlichen Fakultat der Rheinischen Friedrich-Wilhelms-Universitat Bonn. 108 pp.
Grellet-Tinner, 2005. A phylogenetic analysis of oological characters: A case study of saurischian dinosaur relationships and avian evolution. PhD thesis, University of Southern California. 221 pp.
Grellet-Tinner, Chiappe, Norell and Bottjer, 2006. Dinosaur eggs and nesting behaviors: A paleobiological investigation. Palaegeography, Palaeoclimatology, Palaeoecology. 232, 294-321.
Codd, Manning, Norell and Perry, 2007. Avian-like breathing mechanics in maniraptoran dinosaurs. Proceedings of the Royal Society B. 275(1631), 157-161.
Erickson, Rauhut, Zhou, Turner, Inouye, Hu and Norell, 2009. Was dinosaurian physiology inherited by birds? Reconciling slow growth in Archaeopteryx. PLoS ONE. 4(10), e7390.
Nesbitt, Turner, Spaulding, Conrad and Norell, 2009. The theropod furcula. Journal of Morphology. 270, 856-879.
Norton, DML 2000. https://web.archive.org/web/20210603191138/http://dml.cmnh.org/2000Jun/msg00082.html
Balanoff, 2011. Oviraptorosauria: Morphology, phylogeny, and endocranial evolution. PhD thesis. Columbia University. 522 pp.
Balanoff and Norell, 2012. Osteology of Khaan mckennai (Oviraptorosauria: Theropoda). Bulletin of the American Museum of Natural History. 372, 1-77.
Bhullar, Marugan-Lobon, Racimo, Bever, Rowe, Norell and Abzhanov, 2012. Birds have paedomorphic dinosaur skulls. Nature. 487, 223-226.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
Wang, Sun, Sullivan and Xu, 2013. A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of southern China. Zootaxa. 3640(2), 242-257.
Persons, Currie and Norell, 2014 (online 2013). Oviraptorosaur tail forms and functions. Acta Palaeontologica Polonica. 59(3), 553-567.
Pittman and Mallison, 2014. Tail function in oviraptorosaur dinosaurs: Insights from a 3D tail model of Citipati Osmólskae (Theropoda: Oviraptorosauria). Journal of Vertebrate Paleontology. Program and Abstracts 2014, 205-106.
Moyer, Zheng, Norell and Schweitzer, 2015. Microscopic and immunohistochemical analyses of the claw of the nesting dinosaur, Citipati Osmólskae. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 186.
Balanoff, Norell, Hogan and Bever, 2018. The endocranial cavity of oviraptorosaur dinosaurs and the increasingly complex, deep history of the avian brain. Brain, Behavior and Evolution. 91, 125-135.
Norell, Balanoff, Barta and Erickson, 2018. A second specimen of Citipati Osmólskae associated with a nest of eggs from Ukhaa Tolgod, Omnogov Aimag, Mongolia. American Museum Novitates. 3899, 44 pp.
C? sp. nov. (Barsbold, 1981)
Late Campanian, Late Cretaceous
Dzamin Khond, Djadochta Formation, Mongolia
Material- (IGM 100/42; Zamyn Khondt oviraptorid) incomplete skull (180mm), mandibles (153, ~156 mm), fiftteen cervical vertebrae, cervical ribs, seven dorsal vertebrae, three dorsal centra, eighteen dorsal ribs, sacrum, thirty caudal vertebrae (first caudal 32 mm), twenty-three chevrons, scapulacoracoids (scap ~238 mm), furcula, sternal plates (~79, ~84 mm), humeri (205 mm), radius (180 mm), ulna (188 mm), semilunate carpal, metacarpal I (47 mm), phalanx I-1 (95 mm), manual ungual I (73 mm), metacarpal II (104 mm), phalanx II-1 (58 mm), phalanx II-2 (73 mm), manual ungual II (64 mm), metacarpal III (98 mm), phalanx III-1 (42 mm), phalanx III-2 (40 mm), phalanx III-3 (56 mm), manual ungual III (54 mm), ilia (290 mm), pubes (320 mm), ischia, femora (305 mm), tibiae (380 mm), fibula, astragalus, calcaneum, metatarsal I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III (168 mm), phalanx III-1 (39 mm), phalanx III-2, phalanx III-3 (31 mm), pedal ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2 (24 mm), phalanx IV-3 (30.5 mm), phalanx IV-4 (26 mm), pedal ungual IV (45 mm), metatarsal V
Diagnosis- (after Clark et al., 2001) anterior edge of the premaxilla is concave in lateral view; vomers unfused; cervical vertebrae are not elongate.
Comments- This specimen was originally identified as Oviraptor philoceratops (Barsbold, 1981), an identification which persisted unquestioned through the 80's and 90's (e.g. Barsbold, 1983, 1986; Barsbold et al., 1990; Maryanska et al., 2002). The completeness relative to the O. philoceratops holotype resulted in IGM 100/42 becoming the standard example of the species. In 2001, Clark et al. realized the specimen was more similar to the recently named Citipati Osmólskae, and may represent another species of that genus. Clark et al. (2002) reiterated this, while Osmólska et al. (2004) found it to be the sister taxon of Citipati Osmólskae in their cladistic analysis. However, Lu (2004) found that IGM 100/42 was the sister taxon to Oviraptor or Conchoraptor (depending on taxa included) using Maryanska et al.'s (2002) characters. This and the resemblence of the more recently described Nemegtomaia (which was more closely related to Citipati than to IGM 100/42 or Oviraptor in Lu's analysis) suggests we use caution when assigning IGM 100/42 to a particular genus.
The specimen has never been described in detail, though many elements have been illustrated in varied sources. Barsbold (1981) illustrated the skull, mandible, furcula, sternal plates and manual phalanges. He later (1983) illustrated the skull in ventral view, mandible in medial view and scapulocoracoid. Barsbold et al. (1990) illustrated the skull in dorsal view, humerus, complete manus, and metatarsus. Jensen (2008) analyzed the skull for his thesis and presented detailed photographs. The almost complete skeleton is mounted, but publically available photos are too small to be useful. The data matrices of Norell et al. (2001), Maryanska et al. (2002) and Osmólska et al. (2004) are the most useful published sources of information besides the figures. Fanti et al. (2012) and Currie et al. (2016) provide measurements.
References- Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological Expedition Transactions. 15, 28-39.
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 19, 1-120.
Barsbold, 1986. The predatory dinosaurs - Oviraptors. In Vorobyeva (ed.). Herpetologische Untersuchungen in Der Mongolischen Volksrepublik. Academia Nauk SSSR. 210-223.
Barsbold, Maryanska and Osmólska, 1990. Oviraptorosauria. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria. University of California Press. 249-258.
Clark, Norell and Barsbold, 2001. Two new oviraptorids (Theropoda: Oviraptorosauria) from the Late Cretaceous Djadokta Formation, Ukhaa Tolgod. Journal of Vertebrate Paleontology. 21(2), 209-213.
Norell, Clark and Makovicky, 2001. Relationships among Maniraptora: Problems and prospects. In Gauthier and Gall (eds.). New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom. 49-67.
Clark, Norell and Rowe, 2002. Cranial anatomy of Citipati Osmólskae (Theropoda, Oviraptorosauria), and a reinterpretation of the holotype of Oviraptor philoceratops. American Museum Novitates. 3364, 24 pp.
Maryanska, Osmólska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47(1), 97-116.
Lu, 2004. Oviraptorid dinosaurs from southern China. PhD thesis, Southern Methodist University. 249 pp.
Osmólska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria, Second Edition. University of California Press. 165-183.
Jensen, 2008. Beak morphology in oviraptorids, based on extant birds and turtles. Masters Thesis. University of Oslo. 48 pp.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
Currie, Funston and Osmólska, 2016 (online 2015). New specimens of the crested theropod dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica. 61(1), 143-157.

cf. Citipati (Kurzanov, 1987)
Middle-Late Campanian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China
Material- ?(AMNH 6570 in part; paratype of Ornithomimus asiaticus) partial manual ungual ?I (~39 mm) (pers. obs.)
(PIN 2549-100) femur (~160 mm) (Kurzanov, 1987)
Comments- Kurzanov (1987) briefly described and figured "a left avimimid femur from the Upper Cretaceous Iren-Nor locality in China (specimen PIN, no. 2549-100)" (translated), which Currie and Eberth (1993) indicate was found by the Sino-Soviet expedition, which means it was collected between June 14 and July 17 at their localities K (= AMNH locality 141?), L or P. Kurzanov referred it to Avimimidae based on the "accessory condyle" (which is just the lateral condyle being separated from the ectocondylar tuber by a fibular groove as in most theropods), the broad intercondylar flexor groove and similarities of the trochanteric crest. Indeed, he stated "the only not very significant difference is expressed in the fusion of the large and small trochanters, while in Avimimus they are separated by a narrow gap." On the other hand, Osmólska (1996) stated "There is a great resemblance between the femur in [Bagaraatan] ostromi and the femur PIN 2549-100" in that "Both femora have similarly shaped proximal and distal ends, ... well pronounced articular heads and femoral necks, the poorly delimited lesser trochanters, which are as high as the greater, and in the presence of the protuberances on the lateral surface." Additionally, "The distal ends of femora are also similarly shaped in both compared forms" with "distinctive tibiofibular crests ('condylus lateralis' in Kurzanov 1987)." While PIN 2549-100 is similar in shape to both Iren Dabasu Avimimus and Bagaraatan, the latter both have accessory trochanters (usually misidentified as a large, distally placed anterior trochanter in Avimimus) which are absent in PIN 2549-100, Avimimus differs from PIN 2549-100 and Bagaraatan in lacking a distal ectocondylar notch defining the tuber, Bagaraatan differs from PIN 2549-100 and Avimimus in having a narrow flexor groove, and PIN 2549-100 differs from at least Avimimus in having a fourth trochanter reduced to "a slight roughness, located almost under the head of the femur on its medial side" (Kurzanov, 1987) (unknown in Bagaraatan). Currie and Eberth (1993) believed PIN 2549-100 "is probably from a troodontid" and "provisionally referred to Saurornithoides", but Averianov and Sues (2012) concluded it "is probably troodontid but cannot be definitely referred to Saurornithoides" and "should be listed as Troodontidae indet." However, scoring this in Hartman et al.'s maniraptoromorph analysis results in identical scorings to Citipati Osmólskae, with one more step needced to move it sister to Avimimus and two more to move it to Troodontidae (as sister to Linhevenator). Given the stratigraphic and geographic proximity, it is provisionally assigned to cf. Citipati here pending description of the holotype's femur (IGM 100/979 and 1004 have crushed and poorly exposed femora).
The hypodigm of Archaeornithomimus asiaticus includes two collections of largely undescribed and unassociated specimens, AMNH 6570 from Third Asiatic Field site 140 and AMNH 6576 from site 141, discovered between April 22 and May 25 1923. Based on personal examination (July 2009), multiple elements in these collections do not belong to Archaeornithomimus, among which is an ungual in a box of phalanges and calcanea under AMNH 6570. It is moderately curved with a large, proximally placed flexor tubercle and resembles both Sinornithoides' pedal ungual I and Citipati's manual ungual I in its preserved portion. As it is twice the length of Iren Dabasu's troodontid pedal unguals I but scales well to PIN 2549-100, the latter identification is provisionally preferred here.
References- Kurzanov, 1987. Avimimidae and the problem of the origin of birds [in Russian]. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 31, 1-95.
Currie and Eberth, 1993. Palaeontology, sedimentology and palaeoecology of the Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia, People's Republic of China. Cretaceous Research. 14, 127-144.
Osmólska, 1996. An unusual theropod dinosaur from the Late Cretaceous Nemegt Formation of Mongolia. Acta Palaeontologica Polonica. 41, 1-38.
Averianov and Sues, 2012. Correlation of Late Cretaceous continental vertebrate assemblages in middle and central Asia. Journal of Stratigraphy. 36(2), 462-485.

unnamed citipatiine (Dong and Currie, 1995)
Late Campanian, Late Cretaceous
Wulansuhai Formation (= Bayan Mandahu Formation), Mongolia
Material- (IVPP V9608) vertebral fragments, proximal scapula, partial furcula, humerus (168 mm), incomplete radius, incomplete ulna, metacarpal I (32 mm), phalanx I-1 (79 mm), partial manual ungual I, metacarpal II (82+ mm), phalanx II-1 (60 mm), phalanx II-2 (66 mm), manual ungual II, incomplete metacarpal III, phalanx III-1 (38 mm), phalanx III-2 (35 mm), phalanx III-3 (40 mm), manual ungual III, partial femur, partial tibia, phalanx II-2 (33 mm), pedal ungual II (28 mm), phalanx III-1, phalanx III-2 (36 mm), phalanx III-3 (28 mm), pedal ungual III, phalanx IV-1 (30 mm), phalanx IV-2 (27 mm), six eggs, egg fragments
Comments- IVPP V9608 was referred to Oviraptor philoceratops by Dong and Currie (1996), but Longrich et al. (2010) referred it to Oviraptorinae indet. and Hartman et al. (2019) recovered it as an oviraptorine in the Conchoraptor+Citipati clade excluded from Khaan+Conchoraptor. After additional taxa were added to the Hartman et al. matrix, it resolved sister to Citipti and "Anomalipes".
Reference- Dong and Currie, 1995. On the discovery of an oviraptorid skeleton on a nest of eggs. Journal of Vertebrate Paleontology. 15(3), 26A.
Dong and Currie, 1996. On the discovery of an oviraptorid skeleton on a nest of eggs at Bayan Mandahu, Inner Mongolia, People's Republic of China. Canadian Journal of Earth Sciences. 33(4), 631-636.
Longrich, Currie and Dong, 2010. A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of Bayan Mandahu, Inner Mongolia. Palaeontology. 53(5), 945-960.

Rinchenia Barsbold, 1997
R. mongoliensis (Barsbold, 1986) Barsbold, 1997
= Oviraptor mongoliensis Barsbold, 1986
= Citipati mongoliensis (Barsbold, 1986) Paul, 2010
Early Maastrichtian, Late Cretaceous
Altan Uul, Nemegt Formation, Mongolia
Holotype- (IGM 100/32A) skull (156 mm), mandibles (150 mm), atlantal intercentrum, atlantal neuropophyses, axial intercentrum, axis, third-eleventh cervical vertebrae, first-eleventh dorsal vertebrae, gastralia, synsacrum (217 mm), twenty-seven caudal vertebrae, fourteen chevrons, scapulae (~203 mm), coracoid, furcula, partial sternal plate, four sternal ribs, humerus (163 mm), incomplete radius (~103.3 mm), incomplete ulna (~116 mm), phalanx I-1 (~72 mm), proximal manual ungual I, three metacarpal or phalangeal fragments, ilium (225 mm), proximal pubis, proximal ischium, femur (287 mm), incomplete tibia (~370 mm), proximal fibula, metatarsal I (~22 mm), phalanx I-1 (~30 mm), pedal ungual I (~22 mm), distal phalanx II-1, pedal ungual II (~28 mm), proximal and distal metatarsal III, proximal phalanx III-1, phalanx III-2 (~36 mm), phalanx III-3 (23.8 mm), pedal ungual III (~32 mm), proximal and distal metatarsal IV, partial phalanx IV-1, phalanx IV-2 (17 mm), phalanx IV-3 (12 mm), phalanx IV-4 (11.9 mm), pedal ungual IV (33 mm)
Diagnosis- (after Funston et al., 2018) tall, domed cranial crest composed primarily of nasals; frontal taller than anteroposteriorly long; postorbital with vertical frontal process parallel to jugal process; interfingering jugal-quadratojugal contact; ventral process of dentary extending posterior to external mandibular fenestra (also in Citipati); angular not contributing to retroarticular process (also in Citipati); rounded hypapophyses on anterior dorsals; six sacral vertebrae; plate-like distal chevrons; unfused scapulocoracoid with straight posteroventral process of coracoid; deltopectoral crest of humerus ventrally tapering; ilium anterodorsally expanded; preacetabular process of ilium ventrally hooked and pointed; tall but anteroposteriorly restricted brevis fossa; low trochanteric crest of femur; popliteal fossa of femur overhung by medial distal condyle.
Other diagnoses- Barsbold (1986) originally distinguished mongoliensis from Oviraptor philoceratops (including IGM 100/42) based on its smaller size, more robust build, and dome-shaped ridge on the skull roof.
Barsbold (1997) later distinguished Rinchenia from Oviraptor (still including IGM 100/42) based on its "higher dome-like crest that incorporates the parietals", "more lightly built" postcranium and lower cervical vertebrae.
Comments- The holotype was discovered in 1984. Barsbold (1986) originally described this specimen as a new species of Oviraptor, but later (1997) assigned it to a new genus, Rinchenia. While Olshevsky (DML, 1997) declared it a nomen nudum because "the redescription of the species and formal assignment as type species of the new genus is in press", these are not valid reasons according to the ICZN. In particular, it satisfies Article 11, Article 13.1.1 by being "accompanied by a description or definition that states in words characters that are purported to differentiate the taxon" ("about the same size as Oviraptor, but the skull has a higher dome-like crest that incorporates the parietals in addition to the premaxillae, nasals, and frontals. The postcranium is more lightly built, and the cervical vertebrae are rather low in comparison with those of Oviraptor"; contra Funston et al., 2018), and Article 13.3 by being "accompanied by the fixation of a type species in the original publication" via Article 68.3 ("When an author establishes a new nominal genus-group taxon for a single taxonomic species and denotes that species by an available name, the nominal species so named is the type species").
The skull and mandible were first illustrated by Barsbold (1986), the ilium by Barsbold et al. (1990), and the twenty-sixth caudal vertebra by Barsbold et al. (2000). Funston et al. (2018) photographs most elements except the manus and some vertebrae. Snively (2000) studies and illustrates a metatarsus as Rinchenia mongoliensis, but the specimen number (IGM 100/42) shows this is actually Citipati sp. nov.. Funston et al. also provides a brief description and states "a complete anatomical description is forthcoming." A detailed description is present in his 2019 thesis. Fanti et al. (2012) provide measurements.
References- Barsbold, 1986. The predatory dinosaurs - Oviraptors. In Vorobyeva (ed.). Herpetologische Untersuchungen in Der Mongolischen Volksrepublik. Academia Nauk SSSR. 210-223.
Barsbold, Maryanska and Osmólska, 1990. Oviraptorosauria. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria. University of California Press. 249-258.
Barsbold, 1997. Oviraptorosauria. In Currie and Padian (eds.). Encyclopedia of Dinosaurs. 505-509.
Olshevsky, DML 1997. https://web.archive.org/web/20210605031503/http://dml.cmnh.org/1997Sep/msg00863.html
Barsbold, Currie, Myhrvold, Osmólska, Tsogtbaatar and Watabe, 2000. A pygostyle from a non-avian theropod. Nature. 403, 155-156.
Snively, 2000. Functional morphology of the tyrannosaund arctometatarsus. Masters Thesis, University of Calgary. 273 pp.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press. 320 pp.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
Funston, Mendonca, Currie and Barsbold, 2018 (online 2017). Oviraptorosaur anatomy, diversity and ecology in the Nemegt Basin. Palaeogeography, Palaeoclimatology, Palaeoecology. 494, 101-120.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria (Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.

Wulatelong Xu, Tan, Wang, Sullivan, Hone, Han, Ma, Tan and Xiao, 2013
W. gobiensis Xu, Tan, Wang, Sullivan, Hone, Han, Ma, Tan and Xiao, 2013
Late Campanian, Late Cretaceous
Wulansuhai Formation, Inner Mongolia, China
Holotype- (IVPP V18409) (29 kg; adult) incomplete skull, partial mandible, eleven partial dorsal vertebrae, four partial dorsal ribs, first sacral centrum, sixteen partial caudal vertebrae, incomplete scapulocoracoids, sternal plates (one partial, one incomplete), partial humerus, metacarpal I (~35 mm), incomplete phalanx I-1 (78 mm), manual ungual I, metacarpal II (~76 mm), phalanx II-1 (35 mm), phalanx II-2 (33 mm), partial manual ungual II, phalanx III-1 (51 mm), partial phalanx III-2, phalanx III-3 fragment, manual ungual III fragment, incomplete ilium (~225 mm), incomplete pubis (~250 mm), partial ischium, incomplete femur (255 mm), incomplete tibia (325 mm), incomplete fibula, proximal tarsus, distal tarsal IV, incomplete metatarsal II, phalanx II-2 (27 mm), pedal ungual II (45 mm), incomplete metatarsal III (143 mm), phalanx III-1 (43 mm), phalanx III-2 (30 mm), phalanx III-3 (24 mm), pedal ungual III, metatarsal IV (139 mm), phalanx IV-1 (30 mm), phalanx IV-2 (23 mm), phalanx IV-3 (20 mm), phalanx IV-4 (16 mm), pedal ungual IV (35 mm), metatarsal V
Diagnosis- (after Xu et al., 2013) ventral extremity of external naris located below midheight of premaxilla; strap-like jugal process of maxilla extends well beyond preorbital bar posteriorly and overlaps jugal; anterodorsal process of surangular basally constricted in lateral view.
Comments- The holotype was discovered in 2009. This entry switches the identification of manual digits II and III (III and IV of the authors) based on figure 2A, as there is no room for a third non-ungual phalanx in the uppermost digit. The length of the complete phalanx in the other digit and fragmentary nature of more distal phalanges makes this uncertain, however. Xu et al. (2013) proposed Wulatelong was a basal oviraptorid, but it clades with Oviraptor when additional taxa are added to the Hartman et al. analaysis.
Reference- Xu, Tan, Wang, Sullivan, Hone, Han, Ma, Tan and Xiao, 2013. A new oviraptorid from the Upper Cretaceous of Nei Mongol, China, and its stratigraphic implications. Vertebrata PalAsiatica. 51(2), 85-101.

"Tongtianlong" Lu, Chen, Brusatte, Zhu and Shen, 2016
"T. limosus" Lu, Chen, Brusatte, Zhu and Shen, 2016
Late Cretaceous
Nanxiong Group, No. 3 high school of Ganxian, Ganxian District, Ganzhou, Jiangxi, China
Material- (DYM-2013-8) skull (~189 mm), mandible (119 mm), (cervical series 410 mm) atlas, axis (17 mm), third cervical vertebra ( mm), fourth cervical vertebra ( mm), fifth cervical vertebra ( mm), sixth cervical vertebra ( mm), seventh cervical vertebra ( mm), eighth cervical vertebra ( mm), ninth cervical vertebra ( mm), tenth cervical vertebra ( mm), eleventh cervical vertebra ( mm), (dorsal series 350 mm) several partial dorsal vertebrae, dorsal ribs, sacrum, nineteen caudal vertebrae, chevrons, incomplete scapulae (185 mm), coracoids (72 mm), furcula, partial sternum, humeri (131 mm), incomplete radius (102 mm), fragmentary ilium, partial pubes, partial ischia, partial femur (250 mm), incomplete tibia (320 mm), incomplete fibula, astragalus, calcaneum, partial tarsometatarsus (II 125, III 135, IV 110 mm), phalanx II-1, phalanx II-2, pedal ungual II, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal V (38 mm)
Diagnosis- (after Lu et al., 2016) dome-like skull roof with highest point located above posterodorsal corner of orbit; anterior margin of premaxilla highly convex in lateral view; distinct process at middle of anterior parietal margin; lacrimal shaft anteroposteriorly long in lateral view, with flat lateral surface; foramen magnum smaller than occipital condyle (also in Incisivosaurus and Anzu); absence of dentary symphyseal ventral process; absence of posterolateral sternal process.
Comments- "Tongtianlong" was first announced by Lu et al. (2015) was later described by Lu et al. on November 10 2016 as a new taxon of oviraptorid. However, this paper has no mention of ZooBank and as of February 6 2020 "Tongtianlong" lacks an entry on the ZooBank website. Thus according to ICZN Article 8.5.3 (an electronic work must "be registered in the Official Register of Zoological Nomenclature (ZooBank) (see Article 78.2.4) and contain evidence in the work itself that such registration has occurred"), "Tongtianlong limosus" Lu et al., 2016 is a nomen nudum that will only be technically valid pending action on behalf of the authors or ICZN as its journal is not published physically.
Lu et al. (2016) recovered this as a heyuannine sister to Wulatelong+Banji in their analysis using a modified version of Maryanska et al.'s oviraptorosaur matrix. In Hartman et al.'s matrix this clades with Oviraptor and Wulatelong when more taxa are added.
References- Lu, Chen, Kobayshi and Lee, 2015. A new oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 168.
Lu, Chen, Brusatte, Zhu and Shen, 2016. A Late Cretaceous diversification of Asian oviraptorid dinosaurs: Evidence from a new species preserved in an unusual posture. Scientific Reports. 6, 35780.

Oviraptor Osborn, 1924b
= "Fenestrosaurus" Osborn, 1924a
O. philoceratops Osborn, 1924b
= "Fenestrosaurus philoceratops" Osborn, 1924a
Late Campanian, Late Cretaceous
Bayn Dzak, Djadochta Formation, Mongolia
Holotype- (AMNH 6517) incomplete skull (~179 mm), mandibles (195 mm), cervical vertebrae 4-12, dorsal vertebrae 1-7, several dorsal ribs, four uncinate processes, scapula (231 mm), partial coracoid, furcula, sternal ribs, humerus (185 mm), radius (163 mm), ulna (147 mm), phalanx I-1 (79 mm), carpometacarpus (107 mm; mcII 95 mm), phalanx II-1 (54 mm), phalanx II-2 (68 mm), manual ungual II (~47 mm), metacarpal III (99 mm), phalanx III-1 (39 mm), phalanx III-2 (37 mm), phalanx III-3 (28 mm), manual ungual III, partial ilium
Referred- ....(AMNH 6508) ~15 eggs, nest (Osborn, 1924b)
....(AMNH 33092) (embryo or nestling) tibia (58.7 mm), metatarsal III, metatarsal IV (Norell, Balanoff, Barta and Erickson, 2018)
Late Cretaceous?
Mongolia?
?(IGM 100/791) (Balanoff and Norell, 2012)
Other diagnoses- (after Osborn, 1924b) eight craniofacial and mandibular fenestrations [all plesiomorphic- external naris, antorbital fenestra, promaxillary fenestra, orbit, laterotemporal fenestra; supratemporal fenestra; external mandibular fenestra- except "md.f.1" which is probably an eroded surangular fossa]; cranium exceeding facial region in length [also in other oviraptorosaurs]; edentulous [also in other caenagnathoids]; skull extremely abbreviate, orbit and fenestrations exceptionally large [vague but at least applicable to other oviraptorids]; large interclavicle [actually the furcula, also in other oviraptorids]; manus tridactyl [also in other maniraptoromorphs]; metacarpals abbreviated [unclear]; digits irregularly elongated as in the Ornithomimidae [plesiomorphic and not like ornithomimids]; elements of digits not compressed laterally [plesiomorphic, stated different from Tanycolagreus and Chirostenotes].
Comments- The first oviraptorid discovered (on July 3, 1923), it was initially annouced as Fenestrosaurus philoceratops in an article by Osborn (1924a). Osborn describes it as "a small birdlike dinosaur, remains of which were found resting on top of one of the nests of dinosaur eggs; hence the specific name philoceratops, signifying "ceratops lover", "egg-eating or ovivorous. This dinosaur was without teeth." Oviraptor was originally referred to the Ornithomimidae by Osborn (1924b).
Oviraptor was originally hypothesized to be an egg-eater (Osborn, 1924) based on close association with a supposed Protoceratops nest with eggs (AMNH 6508). However, Norell et al. (1994) discovered a brooding oviraptorid specimen which showed the nest and eggs belonged to Oviraptor itself. The diet of Oviraptor is still debated although a lizard preserved in the holotype's body cavity (Norell et al., 1995) suggests it was at least partially carnivorous. Norell et al. (2018) reported and illustrated an embryo or nestling hindlimb associated with the specimen (AMNH 33092) to "be described in detail in another paper."
The holotype was only briefly illustrated and described by Osborn (1924b), but a detailed redescription has not yet appeared. Smith's (1992) attempt made many errors, which were corrected by Clark et al. (2002) in their description of the skull. Makovicky (1995) added information about the preserved vertebrae, though the appendicular remains remain largely ignored. Carrano (1998) listed the femoral length as 282.0 mm, dspite the hindlimb being unpreserved. Norell et al. (2018) included photos of the cervical and pectoral areas.
Osborn (1924b) initially assigned Oviraptor to Ornithomimidae, and even after the naming of Oviraptoridae and discovery of more genera, was not placed in a cladistic context until Barsbold et al. (1990). The latter authors positioned it outside Heyuanninae. This is found in nearly all analyses since, despite the massive increase in taxon number. The remaining unknown is whether Oviraptor is basal to Citipati+Heyuannia, or closer to some taxa with short dentaries.
Not Oviraptor- Barsbold (1976) erected Oviraptoridae for the holotype and referred six individuals including IGM 100/20 and 100/21 to Oviraptor philoceratops. They were later (Barsbold, 1986) made the holotype and paratype of a new genus, Conchoraptor. Osmólska (1976) believed these and additional material (three skulls including ZPAL MgD-I/95 and MgD-I/96 and fragmentary postcrania) belonged to a new species, referring to them as Oviraptor sp.. None of these are referred to Oviraptor currently, with ZPAL MgD-I/95 most often being assigned to Conchoraptor (Maryanska et al., 2002; Kundrat, 2007) or "Ingenia" (Paul, 1998). ZPAL MgD-I/96 is crested and from the Nemegt Formation, so may be Rinchenia or Nemegtomaia. Barsbold (1981) referred IGM 100/42 to O. philoceratops, which resulted in this specimen being the standard example of the species through the 80's and 90's (e.g. Barsbold, 1983; Barsbold, 1986; Barsbold et al., 1990; Maryanska et al., 2002). This was finally shown to be incorrect by Clark et al. (2002), who noted a greater resemblence to Citipati Osmólskae, and assigned it to that genus. Citipati specimens, including IGM 100/979 (Clark et al., 1995) and the holotype IGM 100/978 (Webster, 1996) were referred to Oviraptor based on cranial and manual resemblences to IGM 100/42 until the distinctness of Citipati was recognized by Clark et al. (2001). In addition, Rinchenia (IGM 100/32) was originally described as Oviraptor mongoliensis (Barsbold, 1986) before being renamed unofficially by Barsbold in 1997 and officially by Osmólska et al. in 2004. Paul (1988) lumped "Ingenia" into Oviraptor as O. yanshini, but this would result in almost all oviraptorids being Oviraptor in modern phylogenies. Dong and Currie (1996) referred a fragmentary skeleton (IVPP V9608) to Oviraptor philoceratops, but this may be Citipati instead and was referred to Oviraptorinae indet. by Longrich et al. (2010). Morell (1997) labeled IGM 100/1002 Oviraptor before it was made a paratype of Khaan by Clark et al. (2001). Jensen (2008) calls the tall-crested cast PMO X678 Oviraptor sp., which may be the same taxon as a skull and manus on the Witmer Lab website (Witmer, 2012 online) labeled Oviraptor philoceratops, but how taxon this related to Oviraptor requires further study. Balanoff and Norell (2012) list IGM 100/791 as a specimen of Oviraptor philoceratops, but it is mentioned nowhere else except a similar list in Balanoff's (2011) thesis. Numerous other oviraptorid specimens, have been called Oviraptor, but only the holotype can be properly referred to the genus.
References- Osborn, 1924a. The discovery of an unknown continent. Natural History. 24(2), 133-149.
Osborn, 1924b. Three new Theropoda, Protoceratops zone, central Mongolia. American Museum Novitates. 144, 1-12.
Brown and Schlaikjer, 1940. The structure and relationships of Protoceratops. Annals of the New York academy of Sciences. 40(3), 133-266.
Barsbold, 1976a. O novum pozdnemelovom semeystve melkikh teropod (Oviraptoridae fem. n.) Mongolii. Doklady Akademii Nauk SSSR. 226(3), 685-688.
Barsbold, 1976b. A new Late Cretaceous family of small theropods (Oviraptoridae n. fam.) in Mongolia. Doklady Akademia Nauk SSSR. 226, 221-223.
Osmólska, 1976. New light on skull anatomy and systematic position of Oviraptor. Nature. 262, 683-684.
Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological Expedition Transactions. 15, 28-39. [in Russian]
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 19, 1-120. [in Russian]
Barsbold, 1986. The predatory dinosaurs - Oviraptors. In Vorobyeva (ed.). Herpetologische Untersuchungen in Der Mongolischen Volksrepublik. Academia Nauk SSSR. 210-223.
Paul, 1988. The Predatory Dinosaurs of the World. Simon and Schuster Co., New York. 464 pp.
Barsbold, Maryanska and Osmólska, 1990. Oviraptorosauria. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria. University of California Press. 249-258.
Smith, 1990. Osteology of Oviraptor philoceratops, a possible herbivorous theropod from the Upper Cretaceous of Mongolia. Journal of Vertebrate Paleontology. 3(supplement), 42A.
Smith, 1992. The type specimen of Oviraptor philoceratops, a theropod dinosaur from the Upper Cretaceous of Mongolia. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen. 186, 365-388.
Norell, Clark, Dashzeveg, Barsbold, Chiappe, Davidson, McKenna and Novacek, 1994. A theropod dinosaur embryo, and the affinities of the Flaming Cliffs dinosaur eggs. Science. 266, 779-782.
Norell, Clark, Chiappe, and Dashzeveg, 1995. A nesting dinosaur. Nature. 378, 774-776.
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria (Dinosauria: Theropoda). M.S. thesis, Copenhagen University, Copenhagen, Denmark.
Norell, Dingus and Gaffney, 1995. Discovering Dinosaurs. E.J. Knopf. 225 pp.
Dong and Currie, 1996. On the discovery of an oviraptorid skeleton on a nest of eggs at Bayan Mandahu, Inner Mongolia, People's Republic of China. Canadian Journal of Earth Sciences. 33, 631-636.
Webster, 1996. Dinosaurs of the Gobi. National Geographic. 190(1), 70-89.
Barsbold, 1997. Oviraptorosauria. In Currie and Padian (eds.). Encyclopedia of Dinosaurs. 505-509.
Morell, 1997. The Origin of Birds: the Dinosaur Debate. Audubon. March-April, 36-45.
Carrano, 1998. The evolution of dinosaur locomotion: Functional morphology, biomechanics, and modern analogs. PhD thesis, The University of Chicago. 424 pp.
Clark, Norell and Barsbold, 2001. Two new oviraptorids (Theropoda: Oviraptorosauria) from the Late Cretaceous Djadokta Formation, Ukhaa Tolgod. Journal of Vertebrate Paleontology. 21(2), 209-213.
Clark, Norell and Rowe, 2002. Cranial anatomy of Citipati Osmólskae (Theropoda, Oviraptorosauria), and a reinterpretation of the holotype of Oviraptor philoceratops. American Museum Novitates. 3364, 1-24.
Maryanska, Osmólska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Osmólska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel, Dodson and Osmólska, (eds). The Dinosauria, Second Edition. University of California Press (Berkeley). pp. 165-183.
Jensen, 2008. Beak morphology in oviraptorids, based on extant birds and turtles. Masters Thesis. University of Oslo. 48 pp.
Longrich, Currie and Dong, 2010. A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of Bayan Mandahu, Inner Mongolia. Palaeontology. 53(5), 945-960.
Balanoff, 2011. Oviraptorosauria: Morphology, phylogeny, and endocranial evolution. PhD thesis. Columbia University. 522 pp.
Balanoff and Norell, 2012. Osteology of Khaan mckennai (Oviraptorosauria: Theropoda). Bulletin of the American Museum of Natural History. 372, 1-77.
Witmer, 2012 online. https://people.ohio.edu/witmerl/collections/Theropods/oviraptor.htm
Norell, Balanoff, Barta and Erickson, 2018. A second specimen of Citipati Osmólskae associated with a nest of eggs from Ukhaa Tolgod, Omnogov Aimag, Mongolia. American Museum Novitates. 3899, 44 pp.

Heyuanninae Yun, 2019
Definition- (Heyuannia huangi + Conchoraptor gracilis) (Yun, 2019)
= "Ingeniinae" Barsbold, 1981
Definition- ("Ingenia" yanshini + Conchoraptor gracilis) (Osmólska, Currie and Barsbold, 2004)
Comments- Barsbold (1981) created Ingeniinae to separate his new genus Ingenia from Oviraptor (to which he referred specimens now distinguished as Conchoraptor and Citipati sp.). However, as Taylor (DML 2004) noted, the genus is preoccupied by a tripyloidid nematode (Gerlach, 1957) so any family level taxon based on it must refer to nematodes. A search of the nematode literature suggests that the internal struicture of Tripyloididae reamins unresolved and that no nematode subfamily Ingeniinae has been proposed. Once Barsbold named Conchoraptor and Rinchenia (originally Oviraptor mongoliensis), these were excluded from "Ingeniinae" as well. Barsbold et al. (1990) retain this taxonomy, though their cladogram shows oviraptorines to be paraphyletic to "ingeniines", with Conchoraptor closer to "Ingenia" than to Oviraptor. Maryanska et al. (2002) found Conchoraptor and "Ingenia" to be in a clade exclusive of Citipati and Rinchenia, noting this was equivalent to "Ingeniinae" but poorly supported by data. Osmólska et al. (2004) officially defined the taxon, as "Conchoraptor gracilis, Ingenia yanshini, their most recent common ancestor, and all descendants." More recently, in Funston and Currie (2016) Machairosaurus, Jiangxisaurus, Nemegtomiaia and Heyuannia would also fall under this definition. Adding taxa to Hartman et al.'s matrix would also include Khaan and Gobiraptor.
Yun (2019) noted that the subfamily needed to be renamed, so suggested Heyuanninae as a replacement with the definiotion "the least inclusive clade containing Heyuannia huangi and Conchoraptor gracilis." As Heyuannia is almost always sister to "Ingenia"in oviraptorosaur topologies, the definitions usually cover the same taxa. Although the correct stem would be Heyuanniinae, ICZN Article 29.5 states "If a spelling of a family-group name was not formed in accordance with Article 29.3 but is in prevailing usage, that spelling is to be maintained, whether or not it is the original spelling and whether or not its derivation from the name of the type genus is in accordance with the grammatical procedures in Articles 29.3.1 and 29.3.2." More recently Funston et al. (2020) used Heyuanninae as a stem-based group opposing their new subfamily Citipatiinae although they did not explicitly define it as such. If such a definition is proposed in the future, Oviraptor needs to be an additional external specifier because it cannot be a heyuannine according to the ICZN (it must be an oviraptorine).
References- Gerlach, 1957. Die Nematodenfauna des Sandstrandes an der Küste von Mittelbrasilien (Brasilianische Meerse-Nematoden IV). Mitteilungen aus dem Zoologischen Museum in Berlin. 33(2), 411-459.
Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological Expedition Transactions. 15, 28-39. [in Russian]
Barsbold, Maryanska and Osmólska, 1990. Oviraptorosauria. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria. University of California Press. 249-258.
Clark, Norell and Rowe, 2002. Cranial anatomy of Citipati Osmólskae (Theropoda, Oviraptorosauria), and a reinterpretation of the holotype of Oviraptor philoceratops. American Museum Novitates. 3364, 1-24.
Maryanska, Osmólska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Lu, 2004. Oviraptorid dinosaurs from southern China. PhD thesis, Southern Methodist University. 249 pp.
Osmólska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel, Dodson and Osmólska, (eds.). The Dinosauria, Second Edition. University of California Press. 165-183.
Taylor, DML 2004. https://web.archive.org/web/20200928000123/http://dml.cmnh.org/2004Sep/msg00022.html
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.
Yun, 2019. Heyuanninae clade nov., a replacement name for the oviraptorid subfamily "Ingeniinae" Barsbold, 1981. Zootaxa. 4671(2), 295-296.
Funston, Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020. A new two-fingered dinosaur sheds light on the radiation of Oviraptorosauria. Royal Society Open Science. 7: 201184.

undescribed heyuannine (Maryanska et al., 2002)
Early Maastrichtian, Late Cretaceous
Baruungoyot or Nemegt Formation, Mongolia
Material- (ZPAL MgD-I/106) material including partial skull, mandible and postcranium including cervical vertebrae, dorsal vertebrae, caudal vertebrae, chevrons, partial scapulacoracoid, sternal plate, manual ungual I, manual phalanges III-?, ilium, pubis, ischium, partial femur, metatarsal II, metatarsal III and metatarsal IV
Comments- This crestless specimen was listed as Conchoraptor gracilis by Maryanska et al. (2002) and Fanti et al. (2012), the latter of which provide a few measurements. Balanoff (2011) scored it in her thesis and found it resolved as a heyuannine between Heyuannia huangi and H. yanshini. The specimen number indicates it is either from the Baruungoyot or Nemegt Formation.
References- Maryanska, Osmólska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
Balanoff, 2011. Oviraptorosauria: Morphology, phylogeny, and endocranial evolution. PhD thesis. Columbia University. 522 pp.

undescribed heyuannine (U.S. Attorney's Office Southern District of New York, online 2014)
Late Campanian(?), Late Cretaceous
Khulsan or Red Beds of Khermeen Tsav, Baruungoyot Formation, Mongolia
Material- (CMMD coll.) (upper left) (juvenile) skull, mandibles, hyoid, several cervical vertebrae, twelve dorsal vertebrae, several dorsal ribs, gastralia, first sacral vertebra, posterior sacral or proximal caudal centrum, scapula, furcula(?), humerus, radius, ulna, partial ilia, pubes, ischia, femora, tibiae, fibula, astragali, metatarsal I, phalanx I-1, pedal ungual I, metatarsals II, phalanges II-1, phalanges II-2, pedal unguals II, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsals IV, phalanges IV-1, phalanges IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal V
....(upper right) (juvenile) fragmentary skull, mandibles, six posterior cervical vertebrae, few cervical ribs, five anterior dorsal vertebrae, posterior dorsal vertebral fragments, partial dorsal ribs, two posterior sacral centra, humerus, ulna, partial ilium, pubes, ischial fragment, femora, tibiae, fibulae, astragalus, calcaneum, metatarsals I, pedal ungual I, distal tarsal IV, metatarsals II, phalanges II-1, phalanges II-2, pedal unguals II, metatarsals III, phalanges III-1, phalanges III-2, phalanges III-3, pedal unguals III, metatarsals IV, phalanges IV-1, phalanges IV-2, phalanx IV-3, phalanges IV-4, pedal ungual IV
....(cast UALVP 54983-1; lower left) (juvenile) skull, sclerotic plates, mandibles, hyoid, several cervical vertebrae, nine dorsal vertebrae, several dorsal ribs, two uncinate processes, gastralia, sacrum, ~twenty-six caudal vertebrae (c1 23, c3 19, c4 20, c5 18, c6 18 mm), nine chevrons, scapula, coracoids, partial sternum, humerus, radius, ulna, ilia, pubes, ischia, femur, tibia, fibula, astragalus, calcaneum, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal V
....(cast UALVP 54983-3; lower right) (juvenile) posterior dorsal centrum, five posterior dorsal ribs, gastralia, three sacral centra, first to eighth caudal vertebrae (c4 24, c5 24, c6 24 mm), first to eighth chevrons, ilia, pubes (one distal), ischial fragments, femora (one partial), tibiae (one proximal, one distal), fibulae (one proximal, one distal), astragalus, metatarsals II (one partial), partial metatarsals III, partial metatarsals IV
....(center right) (juvenile) fragmented skull, mandibles, several presacral vertebrae, few dorsal ribs, uncinate process, gastralia, scapulae, partial coracoid, humeri, radii, ulnae, carpals, semilunate carpal, metacarpal I, manual ungual I, metacarpal II, phalanx II-?, metacarpal III, phalanx III-2, phalanx III-3, manual unguals III
.... (cast UALVP 54983-2; upper middle) (juvenile) eight proximal caudal vertebrae (c3 20, c4 19, c5 18, c6 18 mm), distal caudal fragments, eight chevrons
Comments- This oviraptorid bonebed was privately owned until a legal case (U.S. Attorney's Office Southern District of New York, online 2014) where smuggled dinosaurs were being returned to the CMMD in Mongolia, including "a rock matrix containing at least four Oviraptors." Casts are available at e.g. Gaston Design (online 2017) as Conchoraptor from the REd Beds of Khermeen Tsav. One is catalogued as UALVP 54983 (Funston, 2019) where they are listed as being Conchoraptor from Khulsan in the online catalogue. Funston et al. (2015) lists caudal and femoral measurements for two individuals, while Persons et al. (2015) lists caudal measurements for three individuals (as 54983-1, 54983-2 and 54983-3), in both references as Conchoraptor gracilis. Unfortunately, the size of each speciem is so similar that it's unsure which skeletons had first caudal and femoral measurements of 23.9 and 213 mm and 22.9 and 212 mm respectively in Funston et al.'s (2015) table.
References- U.S. Attorney's Office Southern District of New York, online 2014. Manhattan U.S. Attorney Announces Return To Mongolia Of Fossils Of Over 18 Dinosaur Skeletons. July 10.
 Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of Alberta, Canada. Cretaceous Research. 54, 179-187.
Persons, Funston, Currie and Norell, 2015. A possible instance of sexual dimorphism in the tails of two oviraptorosaur dinosaurs. Scientific Reports. 5:9472.
Gaston Design, 2017 online. https://www.gastondesign.com/product/818/
 Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria (Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.

Conchoraptor Barsbold, 1986
C. gracilis Barsbold, 1986
= Citipati gracilis (Barsbold, 1986) Paul, 2010
Late Campanian(?), Late Cretaceous
Red Beds of Khermeen Tsav of Baruungoyot Formation, Mongolia
Holotype- (IGM 100/20) skull (~108 mm), posterior mandible
Paratype?* (see Comments)- (IGM 100/38) metacarpal I (24.6 mm), phalanx I-1 (32.9 mm), manual ungual I (~26 mm), metacarpal II (42.8 mm), distal phalanx II-1, phalanx II-2 (26.3 mm), manual ungual II (~23 mm), metacarpal III (41.4 mm), phalanx III-1 (17.6 mm), phalanx III-2 (~19 mm), phalanx III-3 (~25 mm), manual ungual III (~22 mm)
Referred- (IGM 100/21) mandible (93 mm) (Barsbold, 1976)
(IGM 100/36) material including skull, furcula, ilium (176 mm), pubis (145 mm), femur (182 mm), tibia (215 mm), metatarsal III (102.2 mm), phalanx III-1 (31.1 mm), phalanx III-3 (14.6 mm), phalanx IV-2 (13.5 mm), phalanx IV-3 (9.6 mm), phalanx IV-4 (7.3 mm), pedal ungual IV (18.9 mm) (Barsbold 1981)
(IGM 100/39) material including metacarpal I (~23 mm), phalanx I-1 (41 mm), manual ungual I (~27 mm), phalanx II-2 (~31 mm), manual ungual II (~31 mm), metacarpal III (44 mm), phalanx III-3 (~24 mm), manual ungual III (20 mm) (Maryanska et al., 2002)
(IGM 100/46) material including (?)metatarsal II, (?)metatarsal III, pedal phalanx III-3 (18.2 mm), (?)metatarsal IV, phalanx IV-2 (15.9 mm), phalanx IV-3 (11.5 mm), phalanx IV-4 (11.2 mm), pedal ungual IV (24.9 mm), (?)two pedal unguals, (?)metatarsal V (Maryanska et al., 2002)
(IGM 100/47) materal including ilium (173 mm), femur (195 mm) and tibia (258 mm) (Maryanska et al., 2002)
(IGM 100/1275) partial skeleton including sacrum, twenty-seven caudal vertebrae (first caudal 24.5 mm), pygostyle, femur (212 mm), tibia (250 mm), metatarsal III (107.5) and phalanx III-1 (35.3 mm) (Lu et al., 2013)
(IGM 102/2) four mid caudals, nine mid chevrons (Fanti et al., 2012)
(IGM 102/3) material including partial skull, posterior dorsal fragments, dorsal ribs, uncinate processes, gastralia, last three sacral vertebrae, first to tenth caudal vertebrae, first to ninth chevrons, partial scapula, sterna, humerus, radius (99 mm), ulna (100.1 mm), metacarpals I (24 mm), phalanges I-1 (31 mm), manual unguals I (42 mm), metacarpal II (~44 mm), phalanx II-1 (18.3 mm), phalanx II-2 (15.2 mm), manual ungual II (22 mm), metacarpal III, manual ungual III, posterior ilia (226 mm), pubes, ischium, femora (240 mm), tibiae (277 mm), fibula, astragali, calcanea, distal tarsals III, distal tarsals IV, metatarsal I, phalanges I-1, pedal unguals I, metatarsals II, phalanges II-1, phalanges II-2, pedal unguals II, metatarsals III (121 mm), phalanx III-1 (31 mm), phalanx III-2, phalanges III-3 (17.2 mm), pedal unguals III, metatarsals IV, phalanges IV-1, phalanx IV-2 (14.4 mm), phalanx IV-3 (10.9 mm), phalanges IV-4 (8.7 mm), pedal unguals IV (33 mm), metatarsals V (Fanti et al., 2012)
(IGM MAE 97-212 Block 1B) material including ilium (74.5 mm) and femur (81.2 mm) (Lu et al., 2013)
(IGM MAE 97-212 Block 2B) material including ilium (75 mm), femur (80 mm), metatarsal III (52.2 mm) and phalanx III-1 (14.1 mm) (Lu et al., 2013)
(IGM MAE 97-212 Block 7A) material including ilium (77.7 mm) and femur (82.2 mm) (Lu et al., 2013)
(IGM MAE 97-212 Block 7B) material including femur (87.3 mm) and tibia (110 mm) (Lu et al., 2013)
(ZPAL MgD-I/95) incomplete skull (98 mm), mandibles (86 mm), metatarsus (141 mm) (Osmólska, 1976)
(ZPAL MgD-I/101) skull, postcranial fragments (ZPAL online 2006)
?(private coll.) skull, mandible, several cervical vertebrae, few dorsal fragments, dorsal ribs, sacral fragments, over ten caudal vertebrae, at least eight chevrons, scapulae, partial furcula, coracoid, humerus, radius, ulna, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, metacarpal III, phalanx III-1, distal digits II and III, ilia, pubes, ischia, femora, tibiae, fibulae (one proximal), tarsus, metatarsal II, phalanges II-1, phalanges II-2, pedal ungualsII, metatarsal III, phalanges III-1, phalanges III-2, phalanges III-3, pedal ungual III, metatarsal IV, phalanges IV-1, phalanges IV-2, phalanx IV-3, phalanx IV-4, pedal unguals IV, metatarsal (Witmer, 2012 online 1)
?(private coll.) (1.1 m) skull, mandible, eight cervical vertebrae, cervical ribs, thirteen dorsal vertebrae, dorsal ribs, uncinate processes, over thirty caudal vertebrae, several chevrons, scapulae, sternal plates, sternal ribs, ilia, proximal pubis, proximal ischium, femur, tibia, fibula, proximal tarsus, phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal V (Gaston Designs, 2019 online 2)
Late Campanian, Late Cretaceous
Khulsan, Baruungoyot Formation, Mongolia
(UALVP 54984; cast?) material including first-sixth caudal vertebrae (c2 23, c3 23, c4 23, c5 25, c6 23 mm) and four chevrons (Persons et al., 2015)
(UALVP 54986; cast?) material including first-sixth caudal vertebrae (c1 19, c2 18, c3 18, c4 18, c5 17, c6 17 mm) and four chevrons (Persons et al., 2015)
(UALVP 54987; cast?) material including first-sixth caudal vertebrae (c2 20, c3 23, c4 21, c5 24, c6 23 mm) and three chevrons (Persons et al., 2015)
(UALVP 54988; cast?) material including first-sixth caudal vertebrae (c1 26, c3 22, c4 21, c5 20, c6 19 mm) and three chevrons (Persons et al., 2015)
Late Campanian, Late Cretaceous
Ukhaa Tolgod, Djadochta Formation, Mongolia
 (IGM 100/1203) material including posterior axis, third cervical vertebra, dorsal ribs, uncinate processes, scapula, ilium (158 mm), ischium, femur (154 mm), tibia (184 mm), metatarsal III (83.5 mm) and phalanx III-1 (24 mm) (Balanoff, 2011)
(IGM 100/3006) material including partial skull, proximal caudal vertebrae, chevrons, scapula, coracoid, femur (152.7 mm), metatarsal III (87.5 mm) and phalanx III-1 (24.8 mm) (Balanoff, 2011; braincase described by Balanoff et al., 2014)
Early Maastrichtian, Late Cretaceous
Gurilin Tsav, Nemegt Formation, Mongolia
(IGM uncatalogued) material including incomplete skull, mandibles (Funston, online 2018)
Late Campanian(?) or Early Maastrichtian, Late Cretaceous
Baruungoyot or Nemegt Formation, Mongolia
(ZPAL MgD-I/99) material including scapula and ilium (270 mm) (Fanti et al., 2012)
(ZPAL MgD-I/100) material including partial scapula, humerus (108 mm), phalanx I-1 (18 mm), manual ungual I (19 mm), phalanx II-1 (21 mm), phalanx II-2 (22 mm), manual ungual II (24 mm), ilium (210 mm), partial femur (Maryanska et al., 2002)
Late Cretaceous
Mongolia
(FPDM-V6232) skull, skeleton (Goto, Ichishima and Ji, 2005)
(FPDM-V6234) skull, skeleton (Goto, Ichishima and Ji, 2005)
(IGM 97/212) specimen including femur (250 mm) (Erickson et al., 2009)
(IGM 100/19) (Balanoff and Norell, 2012)
(IGM 100/42; note not the same as the Citipati sp. complete skeleton) specimen including tibia (225 mm), phalanx III-3 (17.4 mm), phalanx IV-2 (18.2 mm), phalanx IV-3 (13.1 mm), phalanx IV-4 (11.9 mm), phalanx IV-5 (25.5 mm) (Fanti et al., 2012)
(IGM 100/97) material including femur (185 mm) and tibia (258 mm) (Fanti et al., 2012)
(IGM 102/1) material including partial tibia, phalanx IV-2 (15.2 mm) (Fanti et al., 2012)
(IGM 110/7) material including skull (94 mm), mandible (87.2 mm) (Fanti et al., 2012)
(IGM 110/10) material including mandible (~55 mm) (Fanti et al., 2012)
(IGM 110/11) material including partial skull (Fanti et al., 2012)
(IGM 110/12) material including skull (~101 mm), partial dentary (Fanti et al., 2012)
(IGM 110/18; actually two individuals, as listed below) material including distal caudal vertebrae, pubis and ischium (Lu, 2004)
----(IGM 110/18a) material including humerus (91.1 mm), radius (87.3 mm), ulna (88.2 mm) (Fanti et al., 2012)
----(IGM 110/18b) material including radius (~78.9 mm), ulna (75.1 mm) (Fanti et al., 2012)
(IGM 110/19) material including thirty-two caudal vertebrae, ilium (~115 mm) (Barsbold et al., 2000)
(IGM 110/20) material including partial skull (Fanti et al., 2012)
(IGM 110/21) material including mandible (68.2 mm), humerus (61.3 mm), radius (~54.6 mm), ulna (55 mm), metacarpal I (22.8 mm), partial metacarpal II, tibia (164 mm) (Fanti et al., 2012)
(IGM 110/22) material including partial skull (Fanti et al., 2012)
(IGM 110/25) material including scapula (Fanti et al., 2012)
(IGM 110/26) material including pedal phalanx III-3 (10.6 mm), phalanx IV-2 (13.9 mm), pedal ungual IV (17.9 mm) (Fanti et al., 2012)
(IGM 110/30) material including metacarpal I (26 mm), phalanx I-1 (38 mm), manual ungual I (~30 mm), phalanx II-1 (~27.5 mm), phalanx II-2 (33 mm), manual ungual II (25 mm) (Fanti et al., 2012)
(PIN coll.) skull (92 mm), mandibles (68 mm) (Maryanska et al., 2002)
(PMO X677; cast) skull, mandibles (Jensen, 2008)
(UALVP 49391; cast) skull (118 mm), mandible (89 mm) (Fanti et al., 2012)
(UALVP 49392; cast) skull (99 mm), mandible (86.6 mm) (Fanti et al., 2012)
(cast) skull, mandibles (Witmer, 2012 online 2)
(cast) skull, mandibles (Witmer, 2012 online 3)
Diagnosis- (after Funston et al., 2018) maxilla with large accessory antorbital fenestra; nasals with three dorsal fenestrae per side; postorbital extending nearly to the bottom of the orbit.
Other diagnoses- (after Barsbold, 1986) unreduced second and third manual digits (plesiomorphic); narrow and straight manual unguals II and III (plesiomorphic).
Comments- The holotype of Conchoraptor was discovered in 1971 and photographed by Barsbold (1976). At the time, it was referred to Oviraptor philoceratops. Additionally, a mandible of IGM 100/21 was photographed, while the morphology of these and four other specimens was briefly described. Barsbold (1977) noted cranial characterics of Khermeen Tsav oviraptorids, probably including IGM 100/20 as well as Heyuannia yanshini material. Barsbold (1981) illustrated a furcula of IGM 100/36, referring it and three other "supposedly young" specimens to O. philoceratops as well. These are probably IGM 100/38, 100/39, 100/46 and 100/47 listed in Maryanska et al. (2002) and may be the four specimens noted by Barsbold (1976). In 1986 Barsbold named Conchoraptor gracilis based on IGM 100/20, illustrating a skull and manus as that specimen number. Funston et al. (2018) also list 100/20 as including the manus, but Fanti et al.'s (2012) measurement table would suggest the manus (with incomplete unguals I and II and partial phalanx II-1) is IGM 100/38. Whether 100/36 was separated from 100/20 after 1986, Fanti et al. incorrectly listed 100/20 measurements as 100/36 or some other explanation is correct is unknown. Barsbold et al. (1990) illustrated the skull in dorsal view and also a metatarsus and two pedal unguals, which may belong to IGM 100/46 since it is the only original specimen given pedal measurements by Fanti et al.. The skull was photographed incorrectly mounted on Heyuannia yanshini postcrania in Psihoyos (1994), as later specifically noted by Funston et al. (2018). This same mount (with a mandible that is neither Conchoraptor's nor yanshini's holotype) was displayed at the Nakasato Dinosaur Center as Ingenia and is now at the IGM. Fanti et al. listed IGM 100/36 among yanshini specimens and incorrectly listed IGM 100/80-D as the holotype skull of Conchoraptor, though Funston et al. (2018) state that holotype skull is labeled IGM 100/80-1 on the IGM mount. Funston et al. (2018) photograph and describe the holotype skull and mandible, further described in Funston's (2019) thesis.
Osmólska (1976) described a skull and mandibles (ZPAL MgD-I/95) as Oviraptor sp.. She also noted an additional skull and postcranial fragments from the same formation, and a fragmentary skull from the Nemegt Formation. She believed they and the IGM specimens noted by Barsbold (1976) belonged to a new species. The Nemegt skull (probably ZPAL MgD-I/96) is crested and thus not Conchoraptor, but the other specimens may be. Possible specimen numbers for the second Khermeen Tsav skull are MgD-I/101 (listed on the ZPAL website as oviraptorid), and MgD-I/100 or MgD-I/106 (listed as Conchoraptor by Maryanska et al., 2002; but not given cranial measurements by Fanti et al., making them less likely candidates). ZPAL MgD-I/95 was illustrated as Oviraptor yanshini by Paul (1988), while Elzanowski (1999) described its palate in detail and called it Oviraptor sp.. Specimen numbers indicate all ZPAL specimens are from the Baruungoyot or Nemegt Formations. Maryanska and Osmólska (1997) stated it was probably Conchoraptor or yanshini, along with a few additional fragmentary skulls (GIN 100/30A and two unnumbered GIN specimens called GIN A and B). Most recently, it was listed as Conchoraptor by Maryanska et al. (2002) and Funston et al. (2018) and had its braincase and cranial pneumaticity described by Kundrat (2007) and Kundrat and Janacek (2007), who referred to it as Conchoraptor. Holtz (1994) has been the only publication since Osmólska (1976) to reference the postcrania, listing the metatarsus of ZPAL MgD-I/95 (as Conchoraptor) in a measurement table.
Fanti et al. (2012) lists MPC-D 102/03 as a specimen of Conchoraptor and provides measurements, although they say "a pair of Ingenia collected at Khermeen Tsav [MPC-D 102/02, MPC-D 102/03] by an earlier "Dinosaurs of the Gobi" expedition." Funston et al. (2016) cites the specimen as "Conchoraptor gracilis (MPC-D 102/3; M. A. Norell pers. comm.)" containing multiple individuals, and his thesis confirms they were found at Khermeen Tsav in 2002. Thus the "Ingenia" pair IGM 110/2 and 110/3 in Fanti et al.'s list are probably different, and indeed 102/3 is about twice the size of 110/3, and Funston's (2019) figures confirm 102/3 matches that number in Fanti et al.'s measurement table. Funston reported online (2018) that he was describing an incomplete skull (IGM uncatalogued) found in 2018 and postcranial skeletion (which is IGM 102/3- Funston, 2019) for his 2019 thesis (7-5-2019 Facebook). The first manual digit seems far more robust than the holotype/paratype, but further evaluation pends on publication of the thesis.
Additional material has also been referred to Conchoraptor. Barsbold et al. (2000) listed a tail (IGM 110/19) as belonging to the taxon, while Lu (2004) noted distal caudal vertebrae and pelvic elements from Conchoraptor specimen IGM 110/18. Besides ZPAL MgD-I/100 and 106, Maryanska et al. (2002) refer a PIN specimen and multiple unnumbered IGM specimens to it. The PIN specimen is quite possibly a skull and mandibles referred to yanshini by Glut (1997) and Witmer (online, 2012 4), as Witmer notes it is from the PIN. It was also photographed in Currie (2001). Osmólska et al. (2004) illustrate a skull as Conchoraptor that appears to be neither the holotype, ZPAL MgD-I/95 or the PIN specimen, and may therefore be one of the other specimens mentioned above. Jensen (2008) analyzed a cast for his thesis (PMO X677) which he referred to Conchoraptor gracilis. Erickson et al. (2009) examined the histology of IGM 97/212, which they referred to Conchoraptor gracilis. Balanoff and Norell (2012) mention many postcranial characters, largely from IGM 100/1203 and 100/3006, noting an in prep. paper. They also list IGM 100/19 as a specimen of Conchoraptor gracilis, but it is mentioned nowhere else except a similar list in Balanoff's (2011) thesis. This thesis states both are from Ukhaa Tolgod in the Djadochta Formation, although Funston et al. (2018) stated 100/3006 is from Khulsan in the Baruungoyot Formation. Cervicals and the ribcage of IGM 100/1203 are photographed in Norell et al. (2018). Fanti et al. and Lu et al. (2013) listed numerous specimens as belonging to this species and provided measurements. Among these is IGM 100/1275, which Balanoff and Norell (2012) mention numerous details of and Persons et al. (2013) describe the tail of. The latter authors note this specimen is one of many partial skeletons found together and catalogued as IGM MAE 97-212. Other specimens such as IGM 100/1203 and 3006 may also belong here, and Balanoff et al. (2014) have described the braincase of IGM 100/3006 based on Balanoff's (2011) thesis.
It is especially common to find crestless privately held oviraptorid specimens being referred to Conchoraptor. One such incomplete skeleton ("Conchoraptor Juvenile in 3-D matrix block" on Gaston Design) is listed as being from the Red Beds of Khermeen Tsav, but has manual proportions more similar to Heyuannia, so is provisionally referred to that taxon here. Another specimen (cast is offered by Gaston Design as "Conchoraptor Adult in matrix 44"" - Gaston Design, 2019 online 2) is more difficult to identify as it doesn't preserve arms, though the constricted third metatarsal is unlike Heyuannia yanshini. Witmer Labs has a cast offered by Gaston Designs as "Conchoraptor Juvenile skeleton in 2-D matrix Block". Witmer (2012 online labs) has two skulls referred to Conchoraptor. Persons et al. (2015) list four Conchoraptor UALVP specimens' proximal caudal measurements, which are likely casts such as these. The UAPVP online catalogue assigns each to Khulsan. A slab of six juvenile skeletons at the CMMD (cast UALVP 54983) has been referred to Conchoraptor in the literature (e.g. Funston et al., 2015), but is here placed as a separate heyuannine.
One notable issue surrounding Conchoraptor is the possibility some referred specimens belong to other taxa, as no referred specimen has ever been assigned to the genus based on shared derived characters with the holotype. For instance, Funston et al. (2018) provided the diagnosis listed above and listed ZPAL MgD-I-95 as referrable, but the promaxillary fenestra size has yet to be shown in the holotype, the number of nasal fossae might be seen in the poor quality photo of Osmólska (1976) and the postorbital extent cannot be measured in that specimen. This is particularily worrisome given the discovery of additional uncrested oviraptorids such as Khaan and Heyuannia, and the possibility young specimens of crested oviraptorids may lack crests (e.g. Yulong, Banji). Although some information can be found in Barsbold (1976), Barsbold et al. (1990), Lu (2004), Osmólska et al. (2004), Funston et al (2018), the specimens examined are not specified (a recent exception is Balanoff and Norell, 2012). Similarly, the matrices of Norell et al. (2001), Maryanska et al. (2002), Lu (2004), Fanti et al. (2012) and Lu et al. (2013) are extremely informative and indicate basically every element is represented in at least one specimen, but obviously don't indicate which specimens were used to code each character. Maryanska et al. do list several specimens they used to code their Conchoraptor OTU, but ironically the holotype was not among them. The measurement tables in Fanti et al. and Currie et al. (2016) will prove useful to refer specimens based on ratios once they are examined.
References- Barsbold, 1976a. O novum pozdnemelovom semeystve melkikh teropod (Oviraptoridae fem. n.) Mongolii. Doklady Akademii Nauk SSSR. 226(3), 685-688.
Barsbold, 1976b. A new Late Cretaceous family of small theropods (Oviraptoridae n. fam.) in Mongolia. Doklady Akademia Nauk SSSR. 226, 221-223.
Osmólska, 1976. New light on skull anatomy and systematic position of Oviraptor. Nature. 262, 683-684.
Barsbold, 1977. Kinetism and peculiarities of the jaw apparatus of oviraptors (Theropoda, Saurischia). Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 4, 34-47.
Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological Expedition Transactions. 15, 28-39.
Barsbold, 1986. The predatory dinosaurs - Oviraptors. In Vorobyeva (ed.). Herpetologische Untersuchungen in Der Mongolischen Volksrepublik. Academia Nauk SSSR. 210-223.
Paul, 1988. The Predatory Dinosaurs of the World. Simon and Schuster. 464 pp.
Barsbold, Maryanska and Osmólska, 1990. Oviraptorosauria. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria. University of California Press. 249-258.
Holtz, 1994. The arctometatarsalian pes, an unusual structure of the metatarsus of Cretaceous Theropoda (Dinosauria: Saurischia). Journal of Vertebrate Paleontology. 14, 480-519.
Psihoyos, 1994. Hunting Dinosaurs. Random House. 288 pp.
Glut, 1997. Dinosaurs, the Encyclopedia: Mcfarland & Company, Inc., Publishers, 1076pp.
Maryanska and Osmólska, 1997. The quadrate of oviraptorid dinosaurs. Acta Palaeontologia Polonica. 42, 377-387.
Elzanowski, 1999. A comparison of the jaw skeleton in theropods and birds, with a description of the palate in the Oviraptoridae. Smithsonian Contributions to Paleobiology. 89, 311-323.
Barsbold, Currie, Myhrvold, Osmólska, Tsogtbaatar and Watabe, 2000. A pygostyle from a non-avian theropod. Nature. 403, 155-156.
Currie, 2001. Theropod dinosaurs from the Cretaceous of Mongolia. In Benton, Shishkin, Unwin and Kurochkin (eds.). The Age of Dinosaurs in Russia and Mongolia. 434-455.
Norell, Clark and Makovicky, 2001. Relationships among Maniraptora: Problems and prospects. In Gauthier and Gall (eds.). New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom. 49-67.
Maryanska, Osmólska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Lu, 2004. Oviraptorid dinosaurs from southern China. PhD thesis, Southern Methodist University. 249 pp.
Goto, Ichishima and Ji, 2005. The Flying Dinosaurs. Catalog of an exhibition held at Fukui Kenritsu Kyōryū Hakubutsukan, July 15-November 3, 2005; also to be held at Asahikawa Kagakukan and at Hamamatsu Kagakukan in 2006. Fukui Kenritsu Kyōryū Hakubutsukan. 118 pp.
ZPAL, online 2006. http://www.paleo.pan.pl/collect.htm#Mon-reptilia
Kundrat, 2007. Avian-like attributes of a virtual brain model of the oviraptorid theropod Conchoraptor gracilis. Naturwissenschaften. 94, 499-504.
Kundrat and Janacek, 2007. Cranial pneumatization and auditory perceptions of the oviraptorid dinosaur Conchoraptor gracilis (Theropoda, Maniraptora) from the Late Cretaceous of Mongolia. Naturwissenschaften. 94(9), 769-778.
Jensen, 2008. Beak morphology in oviraptorids, based on extant birds and turtles. Masters Thesis. University of Oslo. 48 pp.
Erickson, Rauhut, Zhou, Turner, Inouye, Hu and Norell, 2009. Was dinosaurian physiology inherited by birds? Reconciling slow growth in Archaeopteryx. PLoS ONE. 4(10), e7390.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press. 320 pp.
Balanoff, 2011. Oviraptorosauria: Morphology, phylogeny, and endocranial evolution. PhD thesis. Columbia University. 522 pp.
Balanoff and Norell, 2012. Osteology of Khaan mckennai (Oviraptorosauria: Theropoda). Bulletin of the American Museum of Natural History. 372, 1-77.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
Witmer, 2012 online 1. https://people.ohio.edu/witmerl/collections/Theropods/conchoraptor3.htm
Witmer, 2012 online 2. https://people.ohio.edu/witmerl/collections/Theropods/conchoraptor2.htm
Witmer, 2012 online 3. https://people.ohio.edu/witmerl/collections/Theropods/conchoraptor1.htm
Witmer, 2012 online 4. https://people.ohio.edu/witmerl/collections/Theropods/Ingenia.htm
Balanoff, Bever and Norell, 2014. Reconsidering the avian nature of the oviraptorosaur brain (Dinosauria: Theropoda). PLoS ONE. 9(12), e113559.
Persons, Currie and Norell, 2014 (online 2013). Oviraptorosaur tail forms and functions. Acta Palaeontologica Polonica. 59(3), 553-567.
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of Alberta, Canada. Cretaceous Research. 54, 179-187.
Persons, Funston, Currie and Norell, 2015. A possible instance of sexual dimorphism in the tails of two oviraptorosaur dinosaurs. Scientific Reports. 5:9472.
Currie, Funston and Osmólska, 2016 (online 2015). New specimens of the crested theropod dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica. 61(1), 143-157.
Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich, 2016. The first oviraptorosaur (Dinosauria: Theropoda) bonebed: Evidence of gregarious behaviour in a maniraptoran theropod. Scientific Reports. 6:35782. DOI: 10.1038/srep35782
Funston, 2018 online. https://gregfunston.com/2018/10/12/introducing/
Funston, Mendonca, Currie and Barsbold, 2018 (online 2017). Oviraptorosaur anatomy, diversity and ecology in the Nemegt Basin. Palaeogeography, Palaeoclimatology, Palaeoecology. 494, 101-120.
Norell, Balanoff, Barta and Erickson, 2018. A second specimen of Citipati Osmólskae associated with a nest of eggs from Ukhaa Tolgod, Omnogov Aimag, Mongolia. American Museum Novitates. 3899, 44 pp.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria (Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Gaston Design, 2019 online 1. https://www.gastondesign.com/product/conchoraptor-juvenile-skeleton-matrix-block/
Gaston Design, 2019 online 2. https://www.gastondesign.com/product/conchoraptor-adult-matrix-2-d/
Balanoff and Norell, in prep.

Jiangxisaurus Wei, Pu, Xu, Liu and Lu, 2013
J. ganzhouensis Wei, Pu, Xu, Liu and Lu, 2013
Late Cretaceous
Nanxiong Group, Longling Town, Nankang District, Ganzhou, Jiangxi, China
Holotype- (HGM41HIII0421) (subadult) skull (150 mm), mandibles (130 mm), hyoid (60 mm), partial atlas, axis (31 mm), third cervical vertebra (~32 mm), fourth cervical vertebra (33 mm), fifth cervical vertebra (33 mm), sixth cervical vertebra (32 mm), seventh cervical vertebra (28 mm), eighth cervical vertebra (25 mm), three dorsal vertebrae (25, 26, 28 mm), nine partial dorsal ribs, first caudal vertebra (25 mm), second caudal vertebra (25 mm), third caudal vertebra (25 mm), fourth caudal vertebra (25 mm), fifth caudal vertebra (25 mm), sixth caudal vertebra (22 mm), seventh caudal vertebra (20 mm), eighth caudal vertebra (20 mm), ninth caudal vertebra (16 mm), three chevrons, scapula, coracoids, incomplete furcula, partial sternal plates, four sternal ribs, humerus (136 mm), radius (96 mm), ulna (95 mm), scapholunare, semilunate carpal, metacarpal I (25 mm), phalanx I-1 (30 mm), manual ungual I, metacarpal II (45 mm), phalanx II-?, metacarpal III (35 mm), phalanx III-?, ilial fragment, pubic fragment, ischial fragment
Diagnosis- (after Wei et al., 2013) elongated mandible (height 20% of length); weakly downturned mandibular symphysis; surangular with elongate and concave lateral surface; radiohumeral radio ~70%.
Comments- Wei et al. (2013) described this taxon as an oviraptorid of uncertain phylogenetic placement, but Funston and Currie (2016) recovered it as a heyuannine closest to Nemegtomaia and Heyuannia. Hartman et al. (2019) also recovered it as a heyuannine but closest to Conchoraptor. It was discovered prior to November 2012.
References- Wei, Pu, Xu, Liu and Lu, 2013. A new oviraptorid dinosaur (Theropoda: Oviraptorosauria) from the Late Cretaceous of Jiangxi Province, Southern China. Acta Geologica Sinica (English Edition). 87(4), 899-904.
Wei, 2014. A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of Nankang city, Jiangxi Province. Masters thesis, [university]. [pp]
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

Oksoko Funston, Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020
O. avarsan Funston, Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020
Early Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia
Holotype- (IGM 102/110a) (~45 kg, >1 year old juvenile) skull (150 mm), scleral plates, mandible (120 mm), hyoid (55.91 mm), axis, third-fourth cervical vertebrae, posterior dorsal vertebrae, partial dorsal ribs, gastralia, synsacrum, first-seventh caudal vertebrae, proximal chevrons, sternal plates, sternal ribs, radius (92 mm), ulna (92 mm), scapholunare, semilunate carpals, two sesamoids, metacarpals I (23.95, 23.5 mm), phalanges I-1 (32.22, 30.1 mm), manual unguals I (31.2, 31.7 mm), metacarpals II (40.09, 37.35 mm), phalanges II-2 (20.82, 20.96 mm), manual unguals II (18.7, 17.7 mm), metacarpals III (28.25, 23.73 mm), phalanx III-1 (9.07 mm), pubes, ischia, femora (235 mm), tibiae (275 mm), fibulae, astragali (46 mm trans), calcaneum, metatarsals I (26 mm), phalanges I-1, pedal unguals I, metatarsals II (112 mm), phalanges II-1, phalanges II-2, pedal unguals II, metatarsals III (127.5 mm), phalanges III-1, phalanges III-2, phalanges III-3, pedal unguals III, metatarsals IV (119 mm), phalanges IV-1, phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal unguals IV, metatarsal V (38 mm)
Paratypes- (IGM 102/11-A) (~33 kg, ~1 year old juvenile) incomplete skull, posterior mandible, atlantal neurapophyses, incomplete axis, third cervical vertebra, fourth cervical vertebra, seventh to tenth dorsal vertebrae, sacrum (146.9 mm), (caudal series ~435 mm) first (20.6 mm) to twenty-seventh caudal vertebrae, ten chevrons, ilia (216 mm), pubes (226 mm), incomplete ischia (166 mm), femora (one proximal; 210 mm), tibiae (one distal; 259 mm), fibulae (one distal), astragali (40.2 mm trans), calcaneum, metatarsals I (20.3 mm), phalanges I-1, pedal unguals I, distal tarsals III, distal tarsals IV, metatarsals II (102 mm), phalanges II-1, phalanges II-2, pedal unguals II, metatarsals III (117 mm), phalanges III-1, phalanges III-2, phalanges III-3, pedal unguals III, metatarsals IV (112 mm), phalanges IV-1, phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal unguals IV
(IGM 102/11-B) (>33 kg) postorbital, quadratojugal, quadrate
(IGM 102/110b) (~45 kg, ~1 year old juvenile) skull (162 mm), scleral plates, mandible, axis, third-fourth cervical vertebrae, dorsal ribs, sacrum, first-fourth caudal vertebrae, two chevrons, humerus (113 mm), radius (90 mm), ulna (94 mm), metacarpal I (24 mm), phalanx I-1 (31.26 mm), manual ungual I (35.2 mm), metacarpal II (39.71 mm), phalanx II-1 (20.6 mm), phalanx II-2 (15 mm), manual ungual II (18 mm), phalanx III-1 (9.23 mm), pubes, ischia (169 mm), femur (224 mm), tibiae (270 mm), fibula, astragalus (42.6 mm trans), metatarsals I (30.2 mm), phalanges I-1, pedal unguals I, metatarsals II (104 mm), phalanges II-1, phalanges II-2, pedal unguals II, metatarsals III (118 mm), phalanges III-1, phalanges III-2, phalanges III-3, pedal unguals III, metatarsals IV (110 mm), phalanges IV-1, phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal unguals IV, metatarsal V (45 mm)
(IGM 102/110c) (~45 kg) three sacral vertebrae, first-seventh caudal vertebrae, proximal chevrons, partial ilium, distal tibia, astragalocalcaneum
Early Maastrichtian, Late Cretaceous
Bugin Tsav, Nemegt Formation, Mongolia
(IGM 100/33; paratype of "Ingenia" yanshini) (subadult) (atlas-axis 29.4 mm) atlas, axis, third cervical vertebra (22.7 mm), fourth cervical vertebra (24.7 mm), fifth cervical vertebra (26.6 mm), sixth cervical vertebra (27.1 mm), (dorsal series 258.9 mm) first-tenth dorsal vertebrae, synsacrum (162.2 mm), (caudal series ~511 mm) twenty-three caudal vertebrae, chevrons, (scapulocoracoid 205 mm) scapulae, coracoids, sternal plates (~60 mm), furcula, humeri (129.3, 128.7 mm), radii (94.7 mm), ulnae (99 mm), metacarpal I (23.8 mm), phalanges I-1 (27.7 mm), manual unguals I (41.7 mm), metacarpal II (41.7 mm), phalanges II-1 (17.5 mm), phalanges II-2 (14.5 mm), manual ungual II (20 mm), metacarpal III (32 mm), pubes (210 mm), ischia (177 mm), femora (233 mm), tibiae (267 mm), fibula, astragalus (48 mm trans), metatarsals I (29.2 mm), phalanges I-1, pedal unguals I, metatarsals II (111.5 mm), phalanges II-1, phalanges II-2, pedal unguals II, metatarsals III (124.5 mm), phalanges III-1 (34.3 mm), phalanges III-2, phalanges III-3 (17.4 mm), pedal unguals III, metatarsals IV (114.5 mm), phalanges IV-1, phalanges IV-2 (13.9 mm), phalanges IV-3 (9.8 mm), phalanges IV-4 (9.2 mm), pedal unguals IV (30.2 mm), metatarsals V (44 mm)
Early Maastrichtian, Late Cretaceous
Gurilin Tsav, Nemegt Formation, Mongolia
(IGM 102/12; 980808 GT SZK Oviraptr) (~74 kg, >5 year old adult) fourth cervical vertebra (30.9 mm), fifth cervical vertebra (30.4 mm), sixth cervical vertebra (30.5 mm), seventh cervical vertebra (28.9 mm), eighth cervical vertebra (29.8 mm), ninth cervical vertebra (28.1 mm), tenth cervical vertebra (27.8 mm), eleventh cervical neural arch, (dorsal series 293.95 mm) first-tenth dorsal vertebrae, four proximal dorsal ribs, (caudal series ~527 mm) fourth to twenty-seventh caudal vertebrae, eight chevrons, pygostyle, manual ungual I (62 mm), incomplete ilium, incomplete ischum, femur (280 mm), tibia (315 mm), fibulae (one partial), astragalocalcaneum, distal tarsals III, distal tarsal IV, metatarsal II (130 mm), metatarsals III (146.4 mm), phalanx III-1, metatarsal IV (134.2 mm)
Diagnosis- (after Funston et al., 2020) apically thickened, dome-shaped cranial crest composed equally of nasals and frontals; nasal recesses housed in a depression; postorbital with dorsally directed frontal process; cervical vertebrae with large epipophyses; functionally didactyl manus; accessory ridge of brevis fossa of ilium; anteriorly curving pubis; large proximodorsal process of distal tarsal IV.
Comments- Funston et al. (2017) mention "three articulated juvenile skeletons, representing a new species of oviraptorid theropod" confiscated from poachers in December 14 2006. They state the taxon has "a domed cranial crest, a functionally didactyl hand, and a short tail." A brief writeup of the presentation by Watson (2017) photographs the specimens, noting they were found at Bugin Tsav and estimated at 45 kg. Funston et al. (2018) calls the taxon represented by all of these specimens the Guriliin Tsav oviraptorid and stated it "will be described elsewhere (Funston et al., submitted)", which was the Funston et al. in review manuscript cited by Funston (2020) and published as Funston et al. (2020). The latter paper states that the precise locality of discovery within the Nemegt Formation is unknown. The specimens are identified as IGM 102/110a, 110b and 110c in order of completeness.
Discovered in 1974, Barsbold (1981) first mentioned IGM 100/33 as a paratype of "Ingenia" yanshini, then illustrated its furcula and sternal plates in 1983. Barsbold et al. (2000) illustrated the nineteenth caudal vertebra and pelvis. The latter appears more similar to the figure in Barsbold et al. (1990) than to the yanshini holotype, perhaps indicating the 1990 figure was based on IGM 100/33. Funston et al. (2018) noted differences from the yanshini holotype such as only six sacral vertebrae (instead of eight), non-hatchet-shaped caudal transverse processes, unfused sterna, a more reduced metacarpal III and no fibulotarsal contact. They also note that Barsbold (1983) expanded the range of yanshini to include Bugin Tsav (Nemegt) in addition to Khermeen Tsav (Baruungoyot), and that "it is possible that MPC-D 100/33 comes from Bugiin Tsav, but there is no detailed locality data for the specimen." By Funston et al. (2020) it was stated to be from Bugin Tsav without reservation. Funston et al. (2018) stated "the sacral count, enlarged first manual digit, reduced third metacarpal, and lack of fibulocalcaneal contact" are shared between the then unnamed Oksoko and IGM 100/33, and the specimen was referred to that specimen when it was described in Funston et al. (2020).
IGM 102/12 (originally HMNS-IGM field number 980807 GT SZK Oviraptr) is from Gurilin Tsav and was said along with 980807 GT Coy Oviraptr to "belong to a single species, which is similar to the partial skeleton of oviraptorosaurians found at Bugin Tsav in 1994 by our party" (Suzuki and Watabe, 2000). However, 980807 GT Coy Oviraptr has a shallower postacetabular process with greater invagination of the ischial peduncle, so is a different taxon. IGM 102/12 was found in August 7 1998, initially photographed it in situ as "Oviraptorosaurian skeleton in Gurilin Tsav" by Suzuki and Watabe, then was later photographed as figure 13 of Tsogtbaatar and Chinzorig (2010) once prepared. In Funston et al. (2015) IGM 102/12 is listed as cf. Ingenia yanshini and given caudal and femoral measurements. Funston and Currie (2016) mentions IGM 102/12 as Ingenia and state it has "downturned, rounded preacetabular blades." The skeleton is photographed on Funston's blog (Funston, 2018 online). Funston et al. (2017) mention it as "an unpoached, partial adult skeleton from Guriliin Tsav", and it was officially referred to their new genus Oksoko by Funston et al. (2020). The latter state "manual ungual I-2 was recovered in 2018 when the site was revisited."
IGM 102/11 was listed the same way as 102/11 by Funston et al. (2015) and Funston and Currie (2016). Funston et al. (2016) figure its distal metatarsal III as "Ingenia yanshiniBarsbold, 1981 (MPC-D 102/011), provenance unknown: confiscated specimen." It was labeled "an unnamed oviraptorid from the Nemegt Formation of Mongolia" by Funston and Currie (2018), who figured its distal tibiotarsus. The skeleton is photographed on Funston's blog (Funston, 2018 online), who assigned it to the same species as IGM 102/12. This shows it is "the second poached skeleton [of] an even younger individual" confiscated in December 14 2006 mentioned in Funston et al.'s (2017) abstract on Oksoko and was referred to the genus by Funston et al. (2020). They note that "associated with it are the postorbital, quadrate and quadratojugal of a slightly larger individual" which they call IGM 102/11-B.
Funston (2019) describes all of these specimens in detail in his thesis, which also includes more figures than the published description. Funston et al. (2020) added Oksoko to the Maryanska et al.-based oviraptorosaur analysis to find it a heyuannine sister to Jiangxisaurus, then Banji.
References- Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological Expedition Transactions. 15, 28-39.
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 19, 1-120.
Barsbold, Maryanska and Osmólska, 1990. Oviraptorosauria. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria. University of California Press. 249-258.
Barsbold, Currie, Myhrvold, Osmólska, Tsogtbaatar and Watabe, 2000. A pygostyle from a non-avian theropod. Nature. 403, 155-156.
Suzuki and Watabe, 2000. Report on the Japan-Mongolia joint paleontological expedition to the Gobi desert, 1998. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 83-98.
Lu, 2004. Oviraptorid dinosaurs from southern China. PhD thesis, Southern Methodist University. 249 pp.
Tsogtbaatar and Chinzorig, 2010. Fossil specimens prepared in Mongolian Paleontological Center: 2002-2008. Hayashibara Museum of Natural Sciences Research Bulletin. 3, 155-166.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of Alberta, Canada. Cretaceous Research. 54, 179-187.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica Polonica. 61(1), 159-173.
Funston, Currie and Tsogtbaatar, 2017. A new oviraptorid (Dinosauria: Theropoda) provides a rare glimpse into social behaviour in dinosaurs. Journal of Vertebrate Paleontology. Program and Abstracts 2017, 116.
Watson, 2017. Roosting dinosaurs are a fossil first. Nature. 548, 510.
Funston, 2018 online. https://gregfunston.com/2018/09/24/mongolia-monday-travel-log-5/
Funston and Currie, 2018. A small caenagnathid tibia from the Horseshoe Canyon Formation (Maastrichtian): Implications for growth and lifestyle in oviraptorosaurs. Cretaceous Research. 92, 220-230.
Funston, Mendonca, Currie and Barsbold, 2018 (online 2017). Oviraptorosaur anatomy, diversity and ecology in the Nemegt basin. Palaeogeography, Palaeoclimatology, Palaeoecology. 494, 101-120.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria (Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Funston, 2020. Caenagnathids of the Dinosaur Park Formation (Campanian) of Alberta, Canada: Anatomy, osteohistology, taxonomy, and evolution. Vertebrate Anatomy Morphology Palaeontology. 8, 105-153.
Funston, Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020. A new two-fingered dinosaur sheds light on the radiation of Oviraptorosauria. Royal Society Open Science. 7: 201184.

Machairasaurus Longrich, Currie and Dong, 2010
M. leptonychus Longrich, Currie and Dong, 2010
Late Campanian, Late Cretaceous
Wulansuhai Formation (= Bayan Mandahu Formation), Inner Mongolia, China
Holotype- (IVPP V15979) distal radius, distal ulna, scapholunare, semilunate carpal, metacarpal I (24 mm), phalanx I-1 (30 mm), manual ungual I (29 mm), metacarpal II (38 mm), phalanx II-1 (21 mm), phalanx II-2 (22 mm), manual ungual II (24 mm), metacarpal III (34 mm), phalanx III-1 (12 mm), phalanx III-2, phalanx III-3, manual ungual III, manual fragments, several pedal phalangeal fragments, pedal ungual I, pedal ungual IV
Referred- (IVPP V15980) dorsal ribs, caudal vertebrae, chevrons, metacarpal I (32 mm), phalanx I-1 (79 mm), manual ungual I (50 mm), phalanx II-1 (60 mm), phalanx II-2 (66 mm), phalanx III-1 (38 mm), partial tibia, fragmentary metatarsal II, phalanx III-3 (28 mm), fragmentary metatarsal IV, phalanx IV-2 (27 mm) (Longrich, Currie and Dong, 2010)
Diagnosis- (after Longrich et al., 2010) manual unguals I–III elongate and bladelike in lateral view (length of claw approximately 400 per cent the height of the proximal articular surface).
Comments- Discovered in 1988 and 1990, the type specimens of Machairasaurus leptonychus were described by Longrich et al. in 2010 as a new taxon. They used a version of Osmólska et al.'s oviraptorosaur matrix which placed Machairasaurus in Heyuanninae, more derived than Khaan and Conchoraptor but more basal than Heyuannia. This holds for future iterations of that matrix as well, such as Funston and Currie (2016), but after adding taxa to Hartman et al.'s matrix the genus falls out closer to Conchoraptor. Fanti et al. (1012) provide measurements.
References- Longrich, Currie and Dong, 2010. A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of Bayan Mandahu, Inner Mongolia. Palaeontology. 53(5), 945-960.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.

Khaan Clark, Norell and Barsbold, 2001
K. mckennai Clark, Norell and Barsbold, 2001
Late Campanian, Late Cretaceous
Ukhaa Tolgod, Djadochta Formation, Mongolia
Holotype- (IGM 100/1127) (female?) skull (~127 mm), mandible (108.4 mm), hyoids, twelve cervical vertebrae, cervical ribs, several dorsal vertebrae, dorsal ribs, uncinate process, gastralia, sacrum, twenty-seven caudal vertebrae (first caudal 18.3 mm), chevrons, scapulacoracoids (154.1 mm; scap 124.7 mm), furcula, sternal plates (58.8 mm), humeri (110.5, 108.6 mm), radii (~102 mm), ulnae (96.5 mm), scapholunare, semilunate carpal, metacarpal I, phalanx I-1, manual ungual I, metacarpal II (47.3 mm), phalanx II-1, phalanx II-2, manual ungual II, metacarpal III (44.4 mm), phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, ilia (182 mm), pubis, ischium, femur (185 mm), tibia (212 mm), fibula (201 mm), metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV (~95 mm), phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV
Paratypes- (IGM 100/973) skull (118.8 mm), mandibles (104 mm), hyoids, atlas, axis, ilia (193.5, 187.5 mm), pubis (156.8 mm), ischium (133.1 mm), femora (188.8, 199 mm), tibiae (221.4, 224 mm), fibula, astragali, calcanea, distal tarsal III, metatarsals I (26.7, ~23 mm), phalanges I-1, pedal ungual I, metatarsals II (82.8, 89.9 mm), phalanges II-1, phalanges II-2, pedal unguals II, metatarsals III (97.7, 98.8 mm), phalanges III-1, phalanges III-2, phalanges III-3, pedal unguals III, metatarsals IV (87.3, 93.9 mm), phalanges IV-1, phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal unguals IV, metatarsal V (35.8 mm)
(IGM 100/1002) (male?) incomplete skull (113.8 mm), mandible (101.6 mm), hyoids, twelve cervical vertebrae, cervical ribs, ten dorsal vertebrae, dorsal ribs, gastralia, sacrum, seven caudal vertebrae, four chevrons, scapulocoracoids (167 mm; scap 138.6 mm), furcula, sternal plates (55.1 mm), sternal ribs, humerus (117.5 mm), radii (95.3 mm), ulnae (~101.6 mm), scapholunare, semilunate carpal, metacarpal I (24.7 mm), phalanx I-1 (38 mm), manual ungual I (44.5 mm), metacarpal II (50.4 mm), phalanx II-1 (26 mm), phalanx II-2 (28.3 mm), manual ungual II (37 mm), metacarpal III (47.6 mm), phalanx III-1 (16.2 mm), phalanx III-2, phalanx III-3, manual ungual III, ilium (186 mm), pubis (~167 mm), ischium (~124 mm), femur (196 mm), tibia (232 mm), fibula (214 mm), astragalus, distal tarsal IV, metatarsal I (23.3 mm), phalanx I-1, pedal ungual I, metatarsals II (91.2 mm), phalanges II-1, phalanges II-2, pedal unguals II, metatarsals III (105.6 mm), phalanges III-1, phalanges III-2, phalanges III-3 (15.3 mm), pedal unguals III, metatarsals IV (98.3 mm), phalanges IV-1, phalanges IV-2 (14 mm), phalanges IV-3 (11.5 mm), phalanges IV-4 (10.7 mm), pedal unguals IV (31 mm), metatarsal V (35.9 mm)
Referred- (IGM 100/3616) multiple individuals (Norell et al., 2018)
Late Cretaceous?
Mongolia?
?(IGM 100/1253) partial skull (Balanoff, 2011)
Diagnosis- (after Clark et al., 2001) metacarpal III is not expanded proximally and does not contact the distal carpals. Differs from Conchoraptor in that the long axis of the external naris is more horizontally oriented and the nasals are fused.
Comments- IGM 100/973 was discovered in 1993 and first photographed in Novacek et al. (1994). It was photographed in Dashzeveg et al. (1995) as cf. Ingenia, and labeled Ingenia yanshini in Webster (1996). IGM 100/1127 and 100/1002 were discovered in 1995, and the latter was photographed in Morell (1997) and labeled Oviraptor. Clark et al. (2001) made these their new taxon Khaan mckennai. IGM 100/1002 and 100/1127 were found in close proximity to each other, and have been called "Romeo and Juliet." Balanoff and Norell (2012) recently described the taxon in depth and Balanoff et al. (2018) described the endocranium, both based on Balanoff's (2011) thesis. Person et al. (2015) noted the proximal chevrons of IGM 100/1002 were more elongate and distally expanded unlike 100/1127, which they considered plausibly sexual dimorphism with 100/1127 being the female.
Gatesy and Middleton (1997) published hindlimb measurements for an "undescribed oviraptorid (Norell pers. comm.)" which is a Khaan specimen based on nearly identical measurements listed by Dyke and Norell (2005). Balanoff (2011) included the partial skull IGM 100/1253 in her oviraptorosaur analysis and recovered it as either sister to Khaan or to Citipati plus Gigantoraptor, but considered it most likely a Khaan specimen.
References- Novacek, Norell, McKenna and Clark, 1994. Fossils of the Flaming Cliffs. Scientific American. 271(6), 60-69.
Dashzeveg, Novacek, Norell, Clark, Chiappe, Davidson, McKenna, Dingus, Swisher and Perle, 1995. Extraordinary preservation in a new vertebrate assemblage from the Late Cretaceous of Mongolia. Nature. 374, 446-449.
Webster, 1996. Dinosaurs of the Gobi. National Geographic. 190(1), 70-89.
Gatesy and Middleton, 1997. Bipedalism, flight, and the evolution of theropod locomotor diversity. Journal of Vertebrate Paleontology. 17(2), 308-329.
Morell, 1997. The Origin of Birds: the Dinosaur Debate. Audubon. March-April, 36-45.
Clark, Norell and Barsbold, 2001. Two new oviraptorids (Theropoda: Oviraptorosauria) from the Late Cretaceous Djadokta Formation, Ukhaa Tolgod. Journal of Vertebrate Paleontology. 21(2), 209-213.
Dyke and Norell, 2005. Caudipteryx as a non-avialan theropod rather than a flightless bird. Acta Palaeontologica Polonica. 50(1), 101-116.
Balanoff and Norell, 2009. Adult morphology and variation within the oviraptorid Khaan mckennai (Theropoda: Oviraptorosauria). Journal of Vertebrate Paleontology. 29(3), 57A.
Balanoff, 2011. Oviraptorosauria: Morphology, phylogeny, and endocranial evolution. PhD thesis. Columbia University. 522 pp.
Balanoff and Norell, 2012. Osteology of Khaan mckennai (Oviraptorosauria: Theropoda). Bulletin of the American Museum of Natural History. 372, 1-77.
Persons, Funston, Currie and Norell, 2015. A possible instance of sexual dimorphism in the tails of two oviraptorosaur dinosaurs. Scientific Reports. 5:9472.
Balanoff, Norell, Hogan and Bever, 2018. The endocranial cavity of oviraptorosaur dinosaurs and the increasingly complex, deep history of the avian brain. Brain, Behavior and Evolution. 91, 125-135.
Norell, Balanoff, Barta and Erickson, 2018. A second specimen of Citipati Osmólskae associated with a nest of eggs from Ukhaa Tolgod, Omnogov Aimag, Mongolia. American Museum Novitates. 3899, 44 pp.

Nemegtomaia Lu, Tomida, Azuma, Dong and Lee, 2005
= Nemegtia Lu, Tomida, Azuma, Dong and Lee, 2004 preoccupied Szczechura, 1978
N. barsboldi (Lu, Tomida, Azuma, Dong and Lee, 2004) Lu, Tomida, Azuma, Dong and Lee, 2005
= Nemegtia barsboldi Lu, Tomida, Azuma, Dong and Lee, 2004
= Citipati barsboldi (Lu, Tomida, Azuma, Dong and Lee, 2004) Paul, 2010
= Conchoraptor barsboldi (Lu, Tomida, Azuma, Dong and Lee, 2004) Paul, 2016
Early Maastrichtian, Late Cretaceous
Nemegt, Nemegt Formation, Mongolia
Holotype- (IGM 100/2112; = PC1001 2112) skull (179 mm), mandibles (153 mm), partial axis, third cervical vertebra (33 mm), fourth cervical vertebra (29 mm), fifth cervical vertebra (32 mm), sixth cervical vertebra (31 mm), seventh cervical vertebra (31 mm), eighth cervical vertebra (33 mm), ninth cervical vertebra (35 mm), tenth cervical vertebra (37 mm), eleventh cervical vertebra (37 mm), twelfth cervical vertebra (36 mm), thirteenth cervical vertebra (33 mm), most cervical ribs, first dorsal vertebrae (31 mm), partial second dorsal vertebra, sixth dorsal neural arch, seventh dorsal neural arch, eighth dorsal vertebra, ninth dorsal vertebra, tenth dorsal vertebra, sacrum (32, 33, 29, 29, 29, 32, 40, ? mm), first caudal neural arch, second caudal neural arch, proximal scapula, distal humeri, radius, ilia (290 mm), proximal pubes, proximal ischia, proximal femur
Early Maastrichtian, Late Cretaceous
Nemegt Formation?, Mongolia
(IGM 107/16) several fragmentary dorsal ribs, distal radius (~99 mm), distal ulna, semilunate carpals, two carpals, metacarpals I (17.7 mm), phalanges I-1 (24.9 mm), manual unguals I (34 mm), metacarpals II (34 mm), phalanges II-1 (16 mm), phalanges II-2 (13.5 mm), manual unguals II (16 mm), metacarpals III (31 mm), phalanges III-1 (one fragmentary; 9.7 mm), proximal phalanx III-2, ilial fragment, incomplete femora (~222 mm) (Fanti et al., 2012)
Early Maastrichtian, Late Cretaceous
Baruungoyot Formation, Mongolia
Referred- (IGM 107/15) partial skull (172 mm), partial mandibles (152 mm), cervical vertebral fragments, dorsal vertebral fragments, rib fragments, partial scapulae (~185 mm), partial humeri (~152 mm), partial radius (~144 mm), partial ulna, partial phalanx I-1, partial manual ungual I, fragmentary metacarpal II, partial phalanx II-2 (18 mm), incomplete manual ungual II (32 mm), metacarpal III fragment, partial pubes, incomplete femora (~286 mm), partial tibiae (~317 mm), partial fibulae, phalanges I-1, pedal unguals I, phalanx II-1, phalanx II-2, phalanx III-3 (25 mm), phalanx IV-2 (23 mm), phalanx IV-3 (16 mm), phalanx IV-4 (18.8 mm), pedal ungual IV (33 mm), nest, egg fragments (Fanti et al., 2012)
Diagnosis- (after Lu et al., 2004) nasopremaxillary crest; anterior margin of crest nearly vertical, nasal process of premaxilla barely exposed in dorsal view; length of frontal approximately one quarter of that of parietal; presence of prefrontal; process on quadrate bears convex surface projecting into cotyla on medial surface of quadratojugal; mandibular condyles of quadrate situated anteriorly to occipital condyle.
Comments- The holotype was discovered in the summer of 1996 and originally called Ingenia sp. by Lu (1999) and Lu et al. (2002), then Oviraptor sp. in Lu (2004). Lu et al. (2004) described it in detail and named it Nemegtia, though it was renamed by Lu et al. (2005) because Nemegtia is a genus of cypridid ostracod (currently considered a junior synonym of Talycypridea). Mounted skeletons incorporating the Nemegtomaia holotype exist but are misleading, as most of the appendicular and caudal material is faked. Note Fanti et al. (2012) claimed IGM 107/16 was from the Nemegt and Baruungoyot Formations in different parts of their paper, but personal communication (1-11-2017) from Fanti indicates the original collection information is incomplete and the associated sediment leaves Fanti favoring a source in the Nemegt Formation. Originally recovered as sister to Citipati in Lu et al. (2004), it has more recently been found to be sister to Heyuannia (Funston and Currie, 2016; Hartman et al., 2019).
References- Szczechura, 1978. Fresh-water ostracodes from the Nemegt Formation (Upper Cretaceous) of Mongolia. Palaeontologia Polonica. 38, 65-121.
Lu, 1999. New material of Ingenia (Barsbold, 1981) from the Nemegt Formation of southwestern Mongolia and its phylogenetic relationships among Oviraptorosauria. Masters Thesis, Institute of Vertebrate Paleontology and Paleoanthropology. 50 pp.
Lu, 2000. Oviraptorosaurs compared to birds. In Shi and Zhang (eds.). Fifth International Meeting of the Society of Avian Paleontology and Evolution and the Symposium on Jehol Biota. Vertebrata PalAsiatica. 38(suppl.), 18.
Lu, Dong, Azuma, Barsbold and Tomida, 2002. Oviraptorosaurs compared to birds. In Zhou and Zhang (eds.). Proceedings of'the 5th Symposium of the Society of Avian Paleontology and Evolution. 175-189.
Lu, 2004. Oviraptorid dinosaurs from southern China. PhD thesis, Southern Methodist University. 249 pp.
Lu, Tomida, Azuma, Dong and Lee, 2004. New oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Nemegt Formation of southwestern Mongolia. Bulletin of the National Science Museum, Tokyo, Series C. 30, 95-130.
Lu, 2005. Oviraptorid dinosaurs from southern China. Geological Publishing House, Beijing. ISBN 7-116-04368-3. 200 pages + 8 plates.
Lu, Tomida, Azuma, Dong and Lee, 2005. Nemegtomaia gen. nov., a replacement name for the oviraptorosaurian dinosaur Nemegtia Lu et al., 2004, a preoccupied name. Bulletin of the National Science Museum, Tokyo, Series C. 31, 51.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press. 320 pp.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910.
Paul, 2016. The Princeton Field Guide to Dinosaurs 2nd Edition. Princeton University Press. 360 pp.
Funston, Mendonca, Currie and Barsbold, 2018 (online 2017). Oviraptorosaur anatomy, diversity and ecology in the Nemegt Basin. Palaeogeography, Palaeoclimatology, Palaeoecology. 494, 101-120.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247

Heyuannia Lu, 2002
= "Ingenia" Barsbold, 1981 preoccupied Gerlach, 1957
= Ajancingenia Easter, 2013a
Comments- The genus Ingenia is preoccupied by the tripyloidid nematode Ingenia mirabilis, as noted by Taylor (DML, 2004). Barsbold is aware of the situation, but Easter (2013a) did not consult him, proposing his own replacement name Ajancingenia in a publication which largely consisted of copied and reworded information from this website used without my permission. While the initial objection to citing The Theropod Database was editorial, Easter did not object to this, lied about consulting with Barsbold in an online forum and took credit for my work in that same forum. Based on my objection, an erratum was published (Easter, 2013b), and while his name remains technically valid according to the ICZN, I do not support its use due to Easter's unprofessional and unethical actions. Funston et al. (2018) proposed a solution where Ajancingenia was synonymized with Heyuannia, its sister taxon in most analyses. Thus "Ingenia" yanshini becomes Heyuannia yanshini, which is accepted here.
References- Gerlach, 1957. Die Nematodenfauna des Sandstrandes an der Küste von Mittelbrasilien (Brasilianische Meerse-Nematoden IV). Mitteilungen aus dem Zoologischen Museum in Berlin. 33(2), 411-459.
Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological Expedition Transactions. 15, 28-39.
Lu, 2002. A new oviraptorosaurid (Theropoda: Oviraptorosauria) from the Late Cretaceous of southern China. Journal of Vertebrate Paleontology. 22(4), 871-875.
Taylor, DML 2004. https://web.archive.org/web/20200928000123/http://dml.cmnh.org/2004Sep/msg00022.html
Easter, 2013a. A new name for the oviraptorid dinosaur "Ingenia" yanshini (Barsbold, 1981; preoccupied by Gerlach, 1957). Zootaxa. 3737(2), 184-190.
Easter, 2013b. Erratum. Zootaxa. 3750(1), 100.
Funston, Mendonca, Currie and Barsbold, 2018 (online 2017). Oviraptorosaur anatomy, diversity and ecology in the Nemegt Basin. Palaeogeography, Palaeoclimatology, Palaeoecology. 494, 101-120.
H. huangi Lu, 2002
= Citipati huangi (Lu, 2002) Paul, 2010
= Conchoraptor huangi (Lu, 2002) Paul, 2016
Late Campanian-Early Maastrichtian, Late Cretaceous
Dongyuan Formation, Huangsha village, Heyuan City, Guangdong, China
Holotype- (HYMV1-1) partial skull (~150 mm), mandible (~150 mm), hyoid (50 mm), thirteen cervical vertebrae (520 mm; fourth 30 mm; sixth ~45 mm), twelve dorsal vertebrae (320 mm), dorsal ribs (30-160 mm), uncnate processes, first sacral vertebra (27 mm), second sacral vertebra (30 mm), third sacral vertebra (30 mm), fourth sacral vertebra (~30 mm), fifth sacral vertebra (~30 mm), sixth sacral vertebra (30 mm), seventh sacral vertebra (25 mm), eighth sacral vertebra (25 mm), first caudal vertebra (26 mm), second caudal vertebra (26 mm), third caudal vertebra (26 mm), fourth caudal vertebra (26 mm), fifth caudal vertebra (26 mm), sixth caudal vertebra (25 mm), seventh caudal vertebra (25 mm), first chevron (60 mm), second chevron (100 mm), third chevron (100 mm), fourth chevron (90 mm), fifth chevron, sixth chevron, proximal scapula, coracoid fragment, furcula, partial ilia (260 mm), pubes (260 mm), ischia (195 mm), femora (255 mm), tibiae (320 mm), proximal fibula, astragalus, calcaneum, metatarsal I (30 mm), phalanx I-1 (19 mm), pedal ungual I (30 mm), metatarsal II (110 mm), phalanx II-1 (35 mm), phalanx II-2 (20 mm), pedal ungual II (45 mm), metatarsal III (135 mm), phalanx III-1 (37 mm), phalanx III-2 (27 mm), metatarsal IV (120 mm), phalanx IV-1 (25 mm), phalanx IV-2 (20 mm), phalanx IV-3 (10 mm), phalanx IV-4 (10 mm)
Paratypes- (HYMV1-2) two dorsal ribs, gastralia, scapula (~175 mm), incomplete coracoid, furcula, partial sternum, sternal ribs, humerus (130 mm), radius (110 mm), ulna (127 mm), scapholunare, semilunate carpal, metacarpal I (32 mm), phalanx I-1, proximal metacarpal II, partial phalanx II-1, phalanx II-2, proximal metacarpal III
(HYMV1-3) phalanx I-1, manual ungual I, distal metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, distal metacarpal III, phalanx III-1
(HYMV1-4) partial pubis, distal femur, proximal tibia, proximal fibula
(HYMV1-5) incomplete manus
Referred- (HYMV1-6) partial skull, mandibular fragment, posterior cervical vertebrae, anterior dorsal vertebrae, dorsal ribs, uncinate processes, gastralia, last ten caudal vertebrae, partial scapulae (~165 mm), coracoids (50 mm; one partial), incomplete furcula, partial sternum, three partial sternal ribs, humeri (one proximal), partial radius, partial ulna, proximal femur, tibia (Lu, 2004)
(HYMV1-7) last three caudal vertebrae, distal metacarpal I, phalanx I-1, manual ungual I, metacarpal II fragment (Lu, 2004)
(HYMV2-1) humerus (105 mm), incomplete radius, ulna (103 mm), scapholunare, semilunate carpal, pisiform, metacarpal I (29 mm), phalanx I-1 (32 mm), manual ungual I (35 mm), metacarpal II (57 mm), phalanx II-1 (17 mm), phalanx II-2 (16 mm), manual ungual II (21 mm), metacarpal III (52 mm) (Lu, 2004)
(HYMV2-2) partial humerus, partial radius, partial ulna (Lu, 2004)
(HYMV2-3) femur, tibia, fibula, astragalus (Lu, 2004)
(HYMV2-4) pectoral girdle (Lu, 2004)
(HYMV2-5) pelvis, partial hindlimb including metatarsal V fragment (Lu, 2004)
(HYMV2-6) ten mid caudal vertebrae (Lu, 2004)
(HYMV2-7) partial pelvis (Lu, 2004)
(HYMV2-8) radius, ulna, metacarpal I (Lu, 2004)
? bones and eggs (Qiu and Huang, 2001)
Late Campanian(?), Late Cretaceous
Red Beds of Khermeen Tsav of Baruungoyot Formation, Mongolia
Referred- ?(private coll.) (840 mm) skull, mandibles, several cervical vertebrae, dorsal vertebra, dorsal ribs, twenty-six caudal vertebrae, eighteen chevrons, scapulae, coracoids, furcula, humeri, radii, ulna, semilunate carpal, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, metacarpal III, phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, ilia, femora, tibia, fibula, metatarsal I, phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal V (Gaston Design, 2019 online)
Diagnosis- (after Lu, 2002) quadratojugal articular surface of quadrate groove-like; quadrate diverticulum enters bone anterolaterally; increased cervical vertebral count; pneumatic foramina present on cervical neural arches and ribs; decreased dorsal vertebral count; increased sacral vertebral count; proximal end of metacarpal I wraps around metacarpal II in ventral view; pubis as long as ilium.
(after Lu, 2004) angle of the fused scapula and coracoid approximately 145°; ratio of coracoid length to the scapular length approximately 0.35; pubis as long as ischium; ratio of femur length to tibia length 0.8.
Comments- First discovered in the summer of 1999, Lu (2002) briefly described Heyuannia, then described it in more depth including additional specimens in his 2004 thesis (published as a book in 2005). The forelimb and pectoral girdle were described further by Lu et al. (2005), though they incorrectly illustrate HYMV1-6 as 1-4. Qiu and Huang (2001) previously mentioned Ingenia bones and supposed oviraptorid eggs from the same locality, which are therefore probably Heyuannia instead. A nearly complete privately owned skeleton has been cast and widely distributed as Conchoraptor ("Conchoraptor Juvenile in 3-D matrix block" on Gaston Design), but its manual proportions more nearly resemble Heyuannia. This it is provisionally referred to Heyuannia huangi here, though it is supposedly from the Red Beds of Khermeen Tsav like Conchoraptor and Heyuannia yanshini.
Cheng et al. (2008) referred two Macroolithus yaotunensis eggs with embryos (Chimei Museum 41 and NMNS-0015726-F02-embryo-01) from the Nanxiong Group of Jiangxi to Heyuannia huangi "or an oviraptorosaurian of similar kind" based on subarctometatarsaly. Yet this is true in most other oviraptorids as well except for Gigantoraptor, "Tongtianlong" and Heyuannia yanshini. Wiemann et al. (2015a, b) used Cheng et al.'s paper to justify assigning all M. yaotunensis eggs to Heyuannia, including specimens from the Hugang Formation of Henan, the Yuanpu or Pingling Formation of Guangdong and the Nanxiong Group of Jiangxi.
Heyuannia's stratigraphy has been confused, with Lu (2002) initially listing it as being from the Dalangshan Formation based on a 1988 source, but that has more recently been confined to the more southern Sanshui Basin, while Heyuannia is from the Heyuan Basin. Similarly, Lu (2004, 2005) used the revised stratigraphy of Zhang et al. 2005 to equate the Heyuan Basin to the nearby Nanxiong Basin and so assigned Heyuannia to the Zhutian Formation of the Nanxiong Group. More recently the stratigraphy of Fang et al. 2004 has been used where in the Heyuan Basin the Dongyuan Formation is equivalent to the Zhutian and Zhenshui Formations of the Nanxiong Group, assigning Heyuannia to that (e.g. Fang et al., 2009).
References- Qiu and Huang, 2001. Dinosaur fossils from the Heyuan Basin in Guangdong Province, China. In Deng and Wang (eds.). Proceedings of the Eighth Annual Meeting of the Chinese Society of Vertebrate Paleontology. 59-63.
Lu, 2002. A new oviraptorosaurid (Theropoda: Oviraptorosauria) from the Late Cretaceous of southern China. Journal of Vertebrate Paleontology. 22(4), 871-875.
Lu, 2004. Oviraptorid dinosaurs from southern China. PhD thesis, Southern Methodist University. 249 pp.
Lu, 2005. Oviraptorid dinosaurs from Southern China. Geological Publishing House, Beijing. 200 pages + 8 plates.
Lu, Huang and Qiu, 2005. The pectoral girdle and the forelimb of Heyuannia (Dinosauria: Oviraptorosauria). In Carpenter (ed.). The Carnivorous Dinosaurs. Indiana University Press. 256-273.
Cheng, Ji, Wu and Shan, 2008. Oviraptorosaurian eggs (Dinosauria) with embryonic skeletons discovered for the first time in China. Acta Geologica Sinica. 82(6), 1089-1094.
Fang, Yue and Ling, 2009. Review of the past 15 years' research on fossil eggs in China. Acta Geoscientica Sinica. 30(4), 523-542.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press. 320 pp.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, Müller and Sander, 2015a. The blue-green eggs of dinosaurs: How fossil metabolites provide insights into the evolution of bird reproduction. PeerJ PrePrints. DOI: 10.7287/peerj.preprints.1080v1
Wiemann, Yang and Sander, 2015b. The colorful eggs of dinosaurs: How fossil metabolites reveal nesting behavior. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 237.
Paul, 2016. The Princeton Field Guide to Dinosaurs 2nd Edition. Princeton University Press. 360 pp.
Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, Müller and Sander, 2017. Dinosaur origin of egg color: Oviraptors laid blue-green eggs. PeerJ. 5:e3706.
Gaston Design, 2019 online. https://www.gastondesign.com/product/817/
H. yanshini (Barsbold, 1981) Funston, Mendonca, Currie and Barsbold, 2018
= "Ingenia" yanshini Barsbold, 1981
= Oviraptor yanshini (Barsbold, 1981) Paul, 1988
= Ajancingenia yanshini (Barsbold, 1981) Easter, 2013
= Conchoraptor yanshini (Barsbold, 1981) Paul, 2016
Late Campanian(?), Late Cretaceous
Red Beds of Khermeen Tsav of Baruungoyot Formation, Mongolia
Holotype- (IGM 100/30) parietal (lost), braincase (lost), posterior mandible, atlas, axis, third cervical vertebra, fourth cervical vertebra with fused ribs, fifth cervical vertebra, sixth cervical vertebra, seventh cervical vertebra, eighth cervical vertebra, ninth cervical vertebra, fourteen dorsal ribs, sacrum, thirty caudal vertebrae (first caudal 25.3 mm), twelve chevrons, scapulae (~145 mm), coracoids, furcula, sternum (~72 mm), humeri (141 mm), ulnae (116 mm), carpometacarpus (mcI 31.6, mcII 50.3, mcIII 46.5 mm), manual ungual I (49.2 mm), phalanx II-1 (21.3 mm), phalanx II-2 (19.8 mm), manual ungual II (25.8 mm), phalanx III-1 (12.8 mm), phalanx III-2 (11 mm), tibiae (290 mm), fibulae, astragalocalcanea, distal tarsal III, metatarsal I, metatarsal II, metatarsal III (125 mm), metatarsal IV
Paratypes- (IGM 100/31) frontal fragment (lost), parietals (lost), mandible (105 mm), postcrania including eight sacral vertebrae (?- see comments), radii (102.6 mm), metacarpal I (30.3 mm), phalanx I-1 (33.5 mm), manual ungual I (46.9 mm), metacarpal II (46.2 mm), phalanx II-1 (18.7 mm), phalanx II-2 (16.2 mm), manual ungual II (25.7 mm), metacarpal III (44 mm), phalanx III-1 (7.2 mm), phalanx III-2 (8 mm), phalanx III-3 (3 mm), ilia (228 mm), pubes (244 mm), ischia, femora (241 mm), tibia, fibula, metatarsal I, phalanx I-1, pedal ungual I, metatarsal II, phalanges II-1, phalanges II-2, pedal unguals II, metatarsal III (119.7 mm), phalanges III-1, phalanges III-2, phalanges III-3 (18.7 mm), pedal unguals III, metatarsal IV, phalanges IV-1, phalanges IV-2 (14.5 mm), phalanges IV-3 (11.5 mm), phalanges IV-4 (10.6 mm), pedal unguals IV (33.3 mm)
(IGM 100/32) mandible (~120 mm), postcrania including cervical vertebrae, twenty-seven caudal vertebrae, partial scapula, humerus (140 mm), radius (110.5 mm), ulna (115.2 mm), semilunate carpal, phalanx I-1 (34.9 mm), manual ungual I (48.9 mm), metacarpal II (50.5 mm), phalanx II-1 (21.9 mm), phalanx II-2 (18.9 mm), manual ungual II (25.7 mm), metacarpal III (41.6 mm), phalanx III-1 (12.6 mm), ilium (242 mm), pubis (220 mm), femur (254 mm), tibia (294 mm), metatarsus (132 mm), phalanx III-1 (33.1 mm), phalanx III-3 (18.9 mm), phalanx IV-2 (16 mm), phalanx IV-3 (1.6 mm), phalanx IV-4 (11.2 mm), pedal ungual IV (32 mm)
Early Maastrichtian, Late Cretaceous
Nemegt, Nemegt Formation, Mongolia
Referred- ?(PJC.2001.2) partial skeleton including metatarsal I, phalanx I-1, metatarsals II, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-4 (Currie, 2002)
Late Cretaceous(?)
Mongolia(?)
(BHI coll.) skull, mandible (Fanti et al., 2012)
(FDPM-V6240) skull, skeleton (Goto, Ichishima and Ji, 2005)
(IGM 100/34) material including scapula, partial humerus (69 mm), ilium (119 mm), femur (135 mm), tibia (162 mm), metatarsus (75.8 mm), phalanx III-1 (20.6 mm), phalanx III-3 (11.8 mm), phalanx IV-2 (10.8 mm), phalanx IV-3 (6.4 mm), phalanx IV-4 (6.2 mm), pedal ungual IV (18 mm) (Snively, 2000)
(IGM 100/35) material including femur (135 mm), partial tibia and phalanx III-1 (20.4 mm) (Maryanska et al., 2002)
?(IGM 100/80-1; = IGM 100/20?; see comments) material including skull (115.6 mm), eleven cervical vertebrae (Lu, 2004)
(IGM 110/02) (Fanti et al., 2012)
(IGM 110/03) material including humerus (73.5 mm), radius (54.7 mm), ulna (55.3 mm), metacarpal I (15.4 mm), phalanx I-1 (16.8 mm), manual ungual I (19 mm), metacarpal II (23.8 mm), phalanx II-1 (9.5 mm) (Fanti et al., 2012)
(IGM PJC2002.17) skull, mandible (Fanti et al., 2012)
?(IGM coll.) material including skull (147.3 mm), mandible (120 mm), femur (238 mm), tibia (275 mm), metatarsal III (127.5 mm) and phalanx III-1 (34.5 mm) (Lu et al., 2013)
?(IGM coll.) material including skull (162 mm), humerus (113 mm), radius (90 mm), ulna (98 mm), metacarpal II (40.6 mm), femur (220 mm), tibia (265 mm), metatarsal III (118 mm) and phalanx III-1 (34.3 mm) (Lu et al., 2013)
Diagnosis- (after Funston et al., 2018) low ridge on medial side of surangular; eight sacral vertebrae (also in Nemegtomaia); chevrons with distal bulb; ulna with large lateral distal process; manual ungual I longer than metacarpal I.
Other diagnoses- (after Barsbold, 1981) highly reduced, slightly curved and compressed unguals of the second and third fingers.
Comments- Barsbold (1977) described cranial characterics of unspecified Khermeen Tsav oviraptorids, including "Ingenia" paratype IGM 100/31 and probably Conchoraptor material. He illustrated a mandible which was later labeled Ingenia by Barsbold et al. (1990) and is probably IGM 100/31 based on the scale bar and measurement in Fanti et al. (2012). "Ingenia" was first described by Barsbold (1981), who illustrated the manual digits of the holotype and mentioned three other specimens in addition to the holotype (IGM 100/31-33). Barsbold (1983) didn't expand on the description, but did illustrate the holotype's braincase, furcula and sternum. Barsbold (1986) later illustrated the humerus, femur and metatarsus of the holotype. Barsbold et al. (1990) added some further information and illustrated the humerus in different views, radius, ulna, complete manus, pelvis and complete hindlimb missing only metatarsal V, pedal phalanx II-2 and digit I. These may all be from the holotype as well, but the specimen numbers were not listed. Psihoyos (1994) includes a photograph later revealed to be a composite of the holotype, IGM 100/31 and 100/32 (Funston et al., 2018), which is largely complete except for the dorsal series, though the skull of the Conchoraptorholotype is mounted on it. This mount was displayed at the Nakasato Dinosaur Center with a mandible that is neither 100/30, 100/31 or 100/20 (previously online photos). Barsbold et al. (2000) illustrated the twenty-seventh caudal vertebra of IGM 100/32. Lu (2004) noted many anatomical details, including noting some of the elements preserved in paratype specimens. Osmólska et al. (2004) illustrated a scapulocoracoid that is different from the holotype and described several anatomical details. Osmólska (2003, 2004) described the skull roof of IGM 100/31, which she probably incorrectly stated was from the White Beds of Khermeen Tsav, which are slightly younger than the Red Beds (Nemegt instead of Baruungoyot). Funston et al. (2018) is the first published source to identify the IGM mount as a composite of IGM 100/30, 100/31 and 100/32 and provides the sources of all elements. They state the braincase of 100/30 and skull roof of 100/31 are lost. Funston et al. also say "There are eight sacral vertebrae (labeled MPCD 100/30), which appear to be fused with the ilia (labeled MPC-D 100/ 31). It is possible that these elements were artificially connected during the creation of the mount, or that the labels are incorrect." This brings Lu's (2004) statement IGM 100/31 has eight sacral vertebrae into question, as it may have been based on this mounted specimen. Fanti et al. (2012) and Currie et al. (2016) provide measurements of numerous elements.
Snively (2000) studied and illustrated the metatarsi of IGM 100/32 and 100/34 as Ingenia yanshini based on photos and measurements from Currie (PJC 1998 II). Maryanska et al. (2002) lists the latter specimen and IGM 100/35 as I. yanshini, while Lu (2004) notes cervical and cranial characters of IGM 100/80-1. The latter is notable as previously the only yanshini cranial material thought to exist was the braincase of the holotype (and possibly different skull roof of IGM 100/31), whereas Lu's statements imply much of the skull is present in IGM 100/80-1. Confusingly, Fanti et al. list IGM 100/80-D as a skull and the holotype of Conchoraptor, and Funston et al, (2018) state the mounted Conchoraptor holotype skull at the IGM is labeled 100/80-1. So perhaps Lu was mistaken in assigning the skull to yanshini, but he also stated IGM 100/80-1 has eleven cervical vertebrae when the IGM mount only has nine cervicals from the yanshini holotype. Currie (2002) noted a new partial skeleton (PJC.2001.2) from the Nemegt Formation which he referred to cf. Ingenia sp.. It may end up to be referrable to Nomingia, Gobiraptor, Rinchenia or Nemegtomaia which are present in that formation. As with Conchoraptor, no evidence has ever been published defending the placement of paratypes or referred material in yanshini and several specimens have been removed (see below). The measurements and other referred specimens are all based on measurement tables in Fanti et al. (2012), except two IGM specimens Lu et al. (2013) call c.f. Ingenia Confiscated 1 and 2. PJC.2001.2 was referred to cf. Ingenia sp. by Currie (2002).
Not "Ingenia"- "Ingenia" yanshini paratype IGM 100/33 was illustrated by Barsbold (1983) and Barsbold et al. (2000), but was removed by Funston et al. (2018) due to having six sacrals, unfused sterna and no fibulotarsal contact among other characters. Paul (1988) illustrated ZPAL MgD-I/95 as Oviraptor yanshini, a combination no other workers follow as it would result in placing most oviraptorids in Oviraptor given recent phylogenies. ZPAL MgD-I/95 is currently assigned to Conchoraptor. Khaan paratype IGM 100/973 was first photographed in Dashzeveg et al. (1995) as cf. Ingenia, and labeled Ingenia yanshini in Webster (1996). This is the source of the Djadochta Formation listing for cf. Ingenia sp. in Weishampel et al. (2004). A skull and mandibles in the PIN collections are referred to Ingenia yanshini by Glut (1997) and Ingenia sp. by Witmer on his website (Witmer, online 2012). This is possibly the PIN coll. specimen referred to Conchoraptor by Maryanska et al. (2002) and is assigned to that genus here. Maryanska and Osmólska (1997) note ZPAL MgD-I/95 and a few additional fragmentary skulls (GIN 100/30A and two unnumbered GIN specimens called GIN A and B) from the Red Beds of Khermeen Tsav may belong to Ingenia or Conchoraptor. The holotype of Nemegtomaia was originally called Ingenia sp. by Lu (1999) and Lu et al. (2002), which is the source of the Nemegt Formation listing for Ingenia sp. in Weishampel et al. (2004). Qiu and Huang (2001) mentioned Ingenia bones and supposed oviraptorid eggs from the locality Heyuannia was later described from in Guangdong, China. They are thus more probably Heyuannia huangi instead of H. yanshini. IGM 110/11 and 110/12 were listed by Funston et al. (2015) as c.f. Ingenia yanshini, but have been reidentified as the unnamed Guriliin Tsav oviraptorid (Funston, 2018 online).
References- Barsbold, 1977. Kinetism and peculiarities of the jaw apparatus of oviraptors (Theropoda, Saurischia). Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 4, 34–47.
Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological Expedition Transactions. 15, 28-39.
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 19, 1-120.
Barsbold, 1986. The predatory dinosaurs - Oviraptors. In Vorobyeva (ed.). Herpetologische Untersuchungen in Der Mongolischen Volksrepublik. Academia Nauk SSSR. 210-223.
Paul, 1988. The Predatory Dinosaurs of the World. Simon and Schuster Co., New York. 464 pp.
Barsbold, Maryanska and Osmólska, 1990. Oviraptorosauria. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria. University of California Press. 249-258.
Psihoyos, 1994. Hunting Dinosaurs. Random House. 288 pp.
Dashzeveg, Novacek, Norell, Clark, Chiappe, Davidson, McKenna, Dingus, Swisher III and Perle, 1995. Unusual preservation in a new vertebrate assemblage from the Late Cretaceous of Mongolia. Nature. 374, 446-449.
Webster, 1996. Dinosaurs of the Gobi. National Geographic. 190(1), 70-89.
Glut, 1997. Dinosaurs, the Encyclopedia. Mcfarland & Company. 1076 pp.
Maryanska and Osmólska, 1997. The quadrate of oviraptorid dinosaurs. Acta Palaeontologia Polonica. 42, 377-387.
Lu, 1999. New material of Ingenia (Barsbold, 1981) from the Nemegt Formation of southwestern Mongolia and its phylogenetic relationships among Oviraptorosauria. Masters Thesis, Institute of Vertebrate Paleontology and Paleoanthropology. 50 pp.
Barsbold, Currie, Myhrvold, Osmólska, Tsogtbaatar and Watabe, 2000. A pygostyle from a non-avian theropod. Nature. 403, 155-156.
Snively, 2000. Functional morphology of the tyrannosaund arctometatarsus. Masters Thesis, University of Calgary. 273 pp.
Norell, Clark and Makovicky, 2001. Relationships among Maniraptora: Problems and prospects. In Gauthier and Gall (eds.). New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom. 49-67.
Qiu and Huang, 2001. Dinosaur fossils from the Heyuan Basin in Guangdong Province, China. In Deng and Wang (eds.). Proceedings of the Eighth Annual Meeting of the Chinese Society of Vertebrate Paleontology. 59-63.
Currie, 2002. Report on fieldwork in Mongolia, September 2001. Alberta Palaeontological Society, sixth annual symposium. 8-12.
Lu, Dong, Azuma, Barsbold and Tomida, 2002. Oviraptorosaurs compared to birds. In Zhou and Zhang (eds.). Proceedings of'the 5th Symposium of the Society of Avian Paleontology and Evolution. 175- 189.
Maryanska, Osmólska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47(1), 97-116.
Osmólska, 2003. Some aspects of the oviraptorosaur (Dinosauria, Theropoda) braincase. 1st EAVP Meeting. 33.
Lu, 2004. Oviraptorid dinosaurs from southern China. PhD thesis, Southern Methodist University. 249 pp.
Osmólska, 2004. Evidence on relation of brain to endocranial cavity in oviraptorid dinosaurs. Acta Palaeontologica Polonica. 49(2), 321-324.
Weishampel, Barrett, Coria, Le Loeuff, Xu, Zhao, Sahni, Gomani and Noto, 2004. Dinosaur Distribution. In Weishampel, Dodson and Osmólska (eds.). The Dinosauria Second Edition. University of California Press. 861 pp.
Goto, Ichishima and Ji, 2005. The Flying Dinosaurs. Catalog of an exhibition held at Fukui Kenritsu Kyōryū Hakubutsukan, July 15-November 3, 2005; also to be held at Asahikawa Kagakukan and at Hamamatsu Kagakukan in 2006. Fukui Kenritsu Kyōryū Hakubutsukan. 118 pp.
Balanoff and Norell, 2012. Osteology of Khaan mckennai (Oviraptorosauria: Theropoda). Bulletin of the American Museum of Natural History. 372, 1-77.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
Witmer, 2012 online. https://people.ohio.edu/witmerl/collections/Theropods/Ingenia.htm
Easter, 2013. A new name for the oviraptorid dinosaur "Ingenia" yanshini (Barsbold, 1981; preoccupied by Gerlach, 1957). Zootaxa. 3737(2), 184-190.
Lu, Currie, Xu, Zhang, Pu and Jia, 2013. Chicken-sized oviraptorid dinosaurs from central China and their ontogenetic implications. Naturwissenschaften. 100(2), 165-175.
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of Alberta, Canada. Cretaceous Research. 54, 179-187.
Currie, Funston and Osmólska, 2016 (online 2015). New specimens of the crested theropod dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica. 61(1), 143-157.
Paul, 2016. The Princeton Field Guide to Dinosaurs 2nd Edition. Princeton University Press. 360 pp.
Funston, 2018 online. https://gregfunston.com/2018/09/24/mongolia-monday-travel-log-5/
Funston, Mendonca, Currie and Barsbold, 2018 (online 2017). Oviraptorosaur anatomy, diversity and ecology in the Nemegt Basin. Palaeogeography, Palaeoclimatology, Palaeoecology. 494, 101-120.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria (Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.

Gobiraptor Lee, Lee, Chinsamy, Lu, Barsbold and Tsogtbaatar, 2019
= "Gobiraptor" Lee, 2018
G. minutus Lee, Lee, Chinsamy, Lu, Barsbold and Tsogtbaatar, 2019
= "Gobiraptor minutus" Lee, 2018
Early Maastrichtian, Late Cretaceous
Altan Uul III, Nemegt Formation, Mongolia
Holotype- (IGM 102/111) (juvenile) partial skull, incomplete mandibles, last sacral vertebra (25.5 mm), first caudal vertebra (23.1 mm), second caudal vertebra (24.3 mm), seven proximal caudal vertebrae (22.8, 24.2, 23.6, 23.6, 23.2, 21.4 mm), proximal chevron fragments, proximal scapula, coracoid fragment(?), proximal humerus, ilia (one partial, one incomplete), incomplete pubis, partial ischium, femora (195.7 mm), distal tarsal III, distal tarsal IV, metatarsal I (28.8 mm), phalanx I-1 (23.1 mm), pedal ungual I, metatarsal II (108.2 mm), metatarsal III (123 mm), phalanx III-1 (36.3 mm), phalanx III-2 (29 mm), proximal phalanx III-3, metatarsal IV (112.4 mm), phalanx IV-1 (20.5 mm), phalanx IV-2 (18.6 mm), phalanx IV-3 (16.2 mm), phalanx IV-4 (12.2 mm), pedal ungual IV, metatarsal V (4.2 mm)
Diagnosis- (after Lee et al., 2019) flat articular surface for quadratojugal on quadrate; anteroposteriorly elongate dentary anterior to external mandibular fenestra; extremely thickened anterodorsal end of mandibular symphysis with posterior expansion of its dorsal surface; rudimentary lingual triturating shelf on dentary bearing small occlusal foramina; weakly developed lingual ridge on each lingual shelf; anterior end of coronoid wedging into ventral surface of dorsal dentary process.
Comments- Discovered in 2008, Lee first named and described this in his 2018 thesis before officially describing and naming it in 2019. In both papers, it emerged as a heyuannine sister to Jiangxisaurus, Banji and "Tongtianlong". When added to the Hartman et al. matrix it emerges sister to Heyuannia yanshini, so may end up falling under the genus Heyuannia if future studies support this.
References- Lee, 2018. Two new maniraptorans (Dinosauria: Theropoda) from the Nemegt Formation (early Maastrichtian) of Mongolia. Masters Thesis, Seoul National University School of Earth and Environmental Sciences. 280 pp.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria (Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Lee, Lee, Chinsamy, Lu, Barsbold and Tsogtbaatar, 2019. A new baby oviraptorid dinosaur (Dinosauria: Theropoda) from the Upper Cretaceous Nemegt Formation of Mongolia. PLoS ONE. 14(2), e0210867.